Foundational Literature for 

Moving Native Plant Materials 

in Changing Climates

Mary I. Williams

R. Kasten Dumroese

Jeremiah R. Pinto 

Martin F. Jurgensen

Plant Transfer

Guidelines 

and Zones

Plant

Migration

Conservation

and 

Restoration

Climate

Change

United States Department of Agriculture

Forest Rocky Mountain General Technical

Service Research Station Report RMRS-GTR-347 December 2015
All Rocky Mountain Research Station publications are written and produced by U.S. Forest Service employees and are in the public domain and 

available at no cost. Even though U.S. Forest Service publications are not copyrighted, they are formatted according to U.S. Department of Agriculture 

standards and research findings and formatting cannnot be altered in reprints. Altering content or formatting including the cover and title page, is strictly 

prohibited.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 i

Foundational Literature for 

Moving Native Plant Materials

in Changing Climates

Mary I. Williams, R. Kasten Dumroese, Jeremiah R. Pinto, and Martin F. Jurgensen

CITATION

Williams, Mary I.; Dumroese, R. Kasten; Pinto, Jeremiah 

R.; Jurgensen, Martin F. 2015. Foundational literature 

for moving native plant materials in changing climates. 

Gen. Tech. Rep. 347. Fort Collins, CO: U.S. Department 

of Agriculture, Forest Service, Rocky Mountain Research 

Station. 303 p.

ABSTRACT

Seed transfer guidelines and zones are used to manage 

the movement of plant materials, but by the end of the 

century many landscapes across the globe will have climates that are incompatible with current vegetation. The 

mismatch in rates between climate change and plant migration and adaptation will pose significant challenges 

for natural resource managers, especially when scientific 

information often lags behind the demand for management 

actions. We developed a bibliography to reflect the growing interest in assisted migration, the intentional movement of plant 

materials in response to climate change, and to provide a central 

foundation for collaboration in generating research questions, 

conducting studies, transferring and acquiring data, expanding 

studies to key species and geographic regions, and guiding 

native plant transfer in changing climates. The bibliography contains literature through 15 March 2015 on plant 

transfer guidelines as related to climate change, restoration and conservation, and adaptation strategies, such 

as assisted migration. References are arranged in four 

chapters: Climate Change, Conservation and Restoration, 

Migration, and Transfer Guidelines and Zones. Chapters 

are further divided into sections: General (editorials, reviews, summaries, surveys, and trends), Research (common garden studies, genecology research, provenance 

tests, range shifts), Strategies (adaptation options, decision support, frameworks, and policies) and Resources 

(manuals, models, tools, and websites).

Keywords: assisted colonization, assisted migration, bibliography, conservation, forestry, land management, managed relocation, restoration

AUTHORS

Mary I. Williams, Michigan Technological University, 

School of Forest Resources and Environmental Science, 

Houghton, Michigan

R. Kasten Dumroese, USDA Forest Service, Grassland, 

Shrubland, and Desert Ecosystems Program, Rocky 

Mountain Research Station, Moscow, Idaho

Jeremiah R. Pinto, USDA Forest Service, Grassland, 

Shrubland, and Desert Ecosystems Program, Rocky 

Mountain Research Station, Moscow, Idaho

Martin F. Jurgensen, Michigan Technological University, School 

of Forest Resources and Environmental Science, Houghton, 

Michigan

ACKNOWLEDGMENTS

This work was supported through agreement 11-JV-11221632-

130 between the U.S. Forest Service (USFS) and Michigan 

Technological University (MTU). Financial assistance was 

provided by MTU and these USFS entities: the National Center 

for Reforestation, Nurseries; and Genetic Resources; the Rocky 

Mountain Research Station (RMRS); and RMRS’ Grassland, 

Shrubland, and Desert Ecosystems Program. We thank Jim 

Marin Graphics for publication design, Niklaas Dumroese for 

the infographics, and RMRS Publishing Services.

Cover images: Climate change and conservation and restoration, 

Mary I. Williams. Plant migration, UNK. Plant transfer guidelines and zones, USDA Forest Service.
ii USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . iii

Appendix A. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii

Chapter 1. Climate Change

General. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

Research. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

Strategies. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49

Resources. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74

Chapter 2. Conservation and Restoration

General. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78

Research. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84

Strategies. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103

Resources. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118

Chapter 3. Migration

General. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122

Research. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136

Strategies. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193

Resources. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 206

Chapter 4. Transfer Guidelines and Zones

General. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 208

Research. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219

Strategies. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 282

Resources. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 290

Table of Contents
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 iii

from current or static planting zones will be growing in unfavorable conditions by the end of this century. Managers may need 

to assist plant species in their migration to new locations. It is 

difficult, however, to stay up-to-date on the latest research, policies, and terminologies. To alleviate this challenge, we compiled 

a bibliography that is focused on climate change, migration, and 

native plant transfer guidelines. It provides a central location for 

native plant transfer research and reflects the growing interest in 

assisted migration. The bibliography is commensurate with several goals set by the USDA, the Forest Service, and the Rocky 

Mountain Research Station (Appendix A, this section), principally the acquisition of research to “ensure our national forests 

and private working lands are conserved, restored, and made 

more resilient to climate change” (USDA 2014, http://www.

usda.gov/documents/usda-strategic-plan-fy-2014-2018.pdf). 

METHODOLOGY

To identify main features of the native plant transfer literature, we searched BioOne, Google Scholar, Science Citation 

Index, ScienceDirect, and Treesearch without date restrictions. We accessed these databases between 6 January 2012 

and 15 March 2015 using 13 search terms in 252 unique search 

strings (fig. 1).

In North America, tree populations are lagging behind their optimal climate niche (415). In other words, some forests are not 

keeping pace with contemporary changes in climate (540) and 

by the end of this century, many landscapes in the United States 

will have climates that are incompatible with current vegetation 

(497) and similar scenarios are possible for Europe (83). Climate 

projections indicate that trees must migrate more than 3,000 m 

per year to avoid maladaptation, extirpation, and/or extinction, 

a rate far exceeding their observed rates of less than 500 m per 

year (346; 383). The disparity in rates between climate change 

and plant migration and adaptation will pose significant challenges for natural resource managers, requiring them to consider a broad range of management options, including assisted 

migration, defined as the intentional movement of plant materials in response to climate change. 

The practice of restricting native plant movement to environments similar to their source has a long history in natural resource 

management, especially in forestry (596; 851). General transfer 

guidelines developed by the U.S. Department of Agriculture 

(USDA) have since improved for most commercially important 

native tree species and a few other native plants, but seed transfer research will need to factor in climate change. Plant materials 

Introduction

Figure 1. Search terms and strings used to locate references for the bibliography. Terms were searched in combinations using “AND” and “OR” 

statements. Asterisks indicate partial search words for which multiple derivations may be relevant, for example, plant, plants, and planting.

native plant* 

OR native seed* 

OR native plant transfer* 

OR native seed transfer* 

zone* OR provenance* 

OR guideline* 

climate change* 

OR global warming 

OR global change*

assisted migration* 

OR assisted colonization* 

OR managed relocation* 

native plant* 

OR native seed* 

OR native plant transfer* 

OR native seed transfer* 

zone* OR provenance* 

OR guideline* 

climate change*

OR global warming 

OR global change*

assisted migration* 

OR assisted colonization* 

OR managed relocation* 

zone* OR provenance* 

OR guideline*

climate change* 

OR global warming 

OR global change* 

assisted migration* 

OR assisted colonization* 

OR managed relocation* 

native plant* 

OR native seed* 

OR native plant transfer* 

OR native seed transfer* 

zone* OR provenance* 

OR guideline* 

climate change*

OR global warming 

OR global change*

assisted migration* 

OR assisted colonization* 

OR managed relocation* 

AND

AND

AND

AND AND

AND AND

AND AND

AND AND AND
iv USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

We focused on references from North America, but considered work having a global perspective and those providing 

novel approaches regardless of geographic location. The database and cross-reference searches produced 877 references, of 

#. Author(s), Date. Title. Source.

Type: 

Geographic Area: 

Keywords: 

Description:

Link:

location of study

keywords written by the 

author(s) or provided by the 

compilers

ID number and

information

abstract, summary, or 

portions of the introduction 

or conclusion written by the 

author(s) or provided by the 

compilers

book, book section, conference 

paper, dissertation, government 

document, journal article, 

newsletter, presentation, report, 

thesis or web page

internet address of reference

Figure 2. Each reference has a unique identification number and contains the following:

Figure 3. Geographic distribution of references included in this bibliography.

Canada: 112

Mexico: 7

North America: 88

USA: 90

Western USA: 196

Eastern USA: 44

Central USA: 9

Northern

Hemisphere: 3

Global: 220

Total: 877

which the main focus is on plant species and populations. We 

reviewed the author, title, publication, date, and full text as 

needed and organized each reference in EndNote. References 

are indexed by author(s) and include date, geographic area, 

keywords, abstract (summary), and URLs, if available (fig. 2). 

When keywords were not provided in the original reference, 

we, the compilers of this bibliography, added them. When an 

abstract was not written by the author(s) but an introduction 

or conclusion that could serve as an abstract (summary) was, 

the later was used and thusly noted. In some cases when no 

suitable summary was available, we generated one. Any portions (keywords or summary) added by the compilers are so 

noted. Most references are peer-reviewed articles from journals (>70%) and the remaining are government documents, 

books, reports, graduate theses, and grey literature, such as 

newsletters, websites, and presentations. This bibliography is 

not an inclusive list; we recognize that this is a current and 

evolving topic. It is our intent, however, as resources permit, 

for the online version of this bibliography to keep pace with 

the growing body of literature on the topic (http://www.rngr.

net/publications/assisted-migration).

SCOPE

The bibliography contains literature that has bearing on native 

plant transfer and includes references about climate change 

(Chapter 1), conservation and restoration (Chapter 2), migration (Chapter 3), and transfer guidelines and zones (Chapter 4). 

It includes work from as early as 1922 by Göte Turesson (611; 

784) who coined the term genecology—the study of species in 

relation to their environment—that provided a foundation for 

researchers to explore genetic × environmental relationships in 

plants and develop seed transfer guidelines for many tree species 

and populations (596). The primary focus is on native plants, but 
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 v

(421; 433; 501; 539; 540), evolutionary and adaptive responses 

(346; 384; 510), and model projections of future plant distributions (448; 497). Assisted migration is a controversial and popular topic in natural resource science and management laden 

with ethical, ecological, and political concerns (339; 340; 342; 

355; 358; 359; 371; 475). In regards to assisted migration, we 

include reviews (354; 388; 466; 577), justifications (158; 363), 

policies (550; 555), decision frameworks (465; 549; 559), guidelines on implementation (416; 568), and strategies to facilitate 

migration, such as using species and population genetic information to guide movement (382; 446; 494; 558). References 

also address reintroduction (385; 419; 459), and assisted migration and transfer of species and populations in Canada (387; 

454; 458; 525), the U.S. (341; 512; 514; 537), China (451), and 

Europe (381; 431).

Native plant transfer guidelines and zones are located in 

Chapter 4, beginning with references that discuss the development and importance of seed transfer guidelines in sustaining 

ecosystem services (302; 584; 588; 596; 597; 599; 611; 612; 613; 

615; 784). Many (>185) references cover genetic × environmental variation in plant species and populations for tree improvement and reforestation (630; 635; 738; 744; 752; 838), and 

delineating transfer zones for future climates (507; 676; 687; 

757; 799). We provide references on how to select appropriate 

seed sources and determine transfer zones (804; 806; 812; 813; 

817; 821; 824), including seed zone mapping tools with climate 

forecasting (830; 845; 853; 854; 877). Seed transfer guidelines 

and zones including maps are located in the Resources section, 

and most guidelines are for tree species, such as Douglas-fir 

(Pseudotsuga menziesii) and ponderosa pine (Pinus ponderosa) 

(633; 862), but there are provisional guidelines for native plants 

(834). Genetic studies and development of seed transfer guidelines for non-commercial plant species have increased over the 

last 15 years (623; 647; 652; 682; 693; 714; 759; 850; 865).

APPLICATION

Climate change poses a significant challenge for restoring, sustaining, and enhancing natural resources. Even in a time where 

information is easy to access and share, it is difficult to keep 

up-to-date of all the latest developments, especially with respect to climate change research, restoration practices, assisted 

migration, and genetic studies. This bibliography can provide 

a central foundation for scientists, land managers, and policymakers to generate research questions, conduct studies and 

literary syntheses, transfer and acquire data, identify knowledge gaps, and guide native plant transfer in a manner to curtail 

significant social, economic, and ecological losses associated 

with impacts from a rapidly changing climate. This research 

has transfered and will continue to transfer, current information to scientists, land managers, policymakers, and the general 

public about climate change and adaptive strategies pertaining 

to native plant species toward maintaining resilient, functional 

ecosystems across the landscape.

we include some references about other organisms, such as the 

well-cited article by Griffith and others (418) about animal translocations and articles by Parmesan and others on butterfly populations (483) and changes in plant and animal distributions due 

to climate change (102; 481; 482). Most references cover work 

from North America (62%) and a few are from Africa, Asia, 

Australia, Europe, and South America. Of the references from 

North America, 62% concern the U.S., 21% represent Canada, 

and a few (<2%) are from Mexico (fig. 3).

Chapters are divided into sections: General (editorials, reviews, summaries, surveys, and trends), Research (common 

garden studies, genecology research, provenance tests, and 

range shifts), Strategies (adaptation options, decision support, 

frameworks, and policies), and Resources (manuals, models, 

tools, and websites). References are alphabetized by author 

within each section. Users can search for topics and words of interest using the ‘Find’ and/or ‘Advanced Search’ tools in Adobe 

Reader (Ctrl+F, Edit > Find/Advanced Search, or right-click the 

document and choose ‘Find’ from the pop-up menu). In the succeeding paragraphs, we provide a brief overview of each chapter 

noting the more prominent references.

Chapter 1 includes references about climate-induced risks to 

plants such as drought (37) and insects (45), changes in phenology due to rising temperatures (102; 481), widespread tree 

mortality (108; 135), and maladaptation in tree populations 

(117). References also cover historical plant distributions (91; 92; 

131) and novel ecosystems (73; 132). Government reports about 

climate change impacts and risks to forests are also included 

(16), as well as government policies or action plans (168; 171; 

202; 876) and reviews and surveys of adaptation strategies (12; 

21; 26; 121; 151). Modeling programs designed to help understand climate change impacts, such as the System for Assessing 

Vulnerability of Species (SAVS) (3; 206; 207), the Forest Tree 

Genetic Risk Assessment System (ForGRAS) (208; 211; 212), 

MaxENT for species habitat modeling (210), and BioSIM to 

forecast pest events (214) can be found in the Resources section 

of Chapter 1. 

In Chapter 2, we provide references about growing native plants 

(328; 329), selecting the right plant material (223; 298), restoring ecosystems (323; 332; 337), producing and storing seed (240; 

247; 272; 330), and conserving sensitive plant species, such as 

five-needle white pines (241; 283; 300; 319). We include theoretical references, such as those discussing paradigm shifts in 

natural resource conservation and restoration (219; 220; 228; 

229; 292) and the value of genetic research and conservation in 

planning for climate change (226; 227; 233; 266; 291; 305; 322). 

Strategies and programs for seed storage (290; 325), native plant 

materials development (320; 321), and genomics (294; 307; 312) 

are also included in this chapter. 

We list more than 240 references concerning both natural and 

human-assisted migration in Chapter 3. It includes studies 

about past vegetation (341; 427; 538), range shifts (423; 449; 

483), migration rates (345; 365; 428; 453), migration capacity 
vi USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

•	 Is a source of technical information for maintaining

and sustaining forestry and agricultural markets and 

public space

•	 Provides information that can be used to develop partnerships and educational material

3 – Apply Knowledge Globally 

•	 Serves as a literary resource to identify needs and set

priorities for forests, grasslands, and shrublands

•	 Exists as a public document and is available on the

Internet

•	 Includes tools for evaluating management practices and

provides appropriate information for decisionmakers

U.S. Forest Service Research and 

Development Research Topics 

http://www.fs.fed.us/research/research-topics/

1 – Invasive Species

•	 Includes research that can be used to help reduce and

minimize the introduction of invasive species in management options, such as assisted migration

•	 Contains literature about the impact of insects and

plant pathogens on forests and wood products

2 – Inventory, Monitoring and Analysis

•	 Provides data, reports, maps, and other resources to

land owners and managers, policy makers, researchers, 

practitioners, and other interest groups

•	 Includes research on the current status and trends of

forests, grasslands, and shrublands, including threats 

and management options

3 – Outdoor Recreation

•	 Provides reports about forest systems and societal benefits

•	 Contains management options to sustain ecosystem

services in a changing climate, such as assisted migration

4 – Resource Management and Use

•	 Provides research and technologies to help sustain the

ecological and economical value of forests, grasslands, 

and shrublands 

•	 Contains information that can be used by communities,

land owners, and practitioners to maintain resources 

and plan for climate change

6 – Wildland Fire and Fuel 

•	 Includes technical guidelines for restoring lands after

wildfire and other disturbances

•	 Provides frameworks for selecting appropriate

plant materials 

Appendix A 

USDA, Forest Service, and Rocky Mountain Research Station 

strategic goals, elements, and research priorities with bulleted 

points showing how the bibliography supports this work.

USDA Strategic Plan, Goals 2014–2018

http://www.usda.gov/documents/usda-strategic-planfy-2014-2018.pdf

1 – Assist rural communities to create prosperity so they are self-sustaining, repopulating, and economically thriving

•	 Is a source of practical forestry and agricultural guidelines that can be used to improve and expand ecosystem

services and products for rural economies

•	 Provides technical information to support current and

future forestry and agricultural markets that are important to rural communities

2 – Ensure national forests and private working lands are conserved, restored, and 

made more resilient to climate change, 

while enhancing our water resources

•	 Is a source of past and current ecological research that

can be used to inform climate change adaptation strategies, such 

as assisted migration

•	 Includes research that can be used to evaluate forest,

grassland, and shrubland health 

U.S. Forest Service Strategic Plan, 

Goals 2015–2020 

http://www.fs.fed.us/strategicplan/

1 – Sustain Our Nation’s Forests and Grasslands

•	 Serves as an integration of work aimed at improving

health and resilience of forests, grasslands, and shrublands

•	 Provides technical information, such as vulnerability

assessment reports, that can be used to foster resilient 

and adaptive ecosystems to mitigate climate change

•	 Includes research on the conservation and restoration

of native plant species, including species with economic 

value

•	 Includes methods, technologies, and strategies for

addressing disturbance

2 – Deliver Benefits to the Public 

•	 Provides information that can be used by the public to

make decisions about managing forests, grasslands, and 

shrublands 
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 vii

Rocky Mountain Research Station 

Strategic Research Priorities 2014–2017 

http://www.fs.fed.us/rmrs/docs/leadership-team/strategicframework.pdf

1 – Disturbance Ecology

•	 Includes research that targets plant response to climate

change and how to mitigate those responses

•	 Is a source of tools to assess, characterize, and model

ecological effects of past, present, and future climate 

change on plant species

•	 Contains research to help understand the effects of

land use and management actions on ecosystems, with 

special emphasis on native plant species

3 – Human-Landscape Interactions

•	 Includes management options and strategies to sustain

ecosystem services in a changing climate

•	 Contains literature that explores social and economic

drivers behind ecosystem change and services 

4 – Inventory and Monitoring

•	 Includes several studies on plant adaptation and migration

•	 Contains new approaches and technologies for monitoring and analyzing ecosystems and landscapes, such 

as plant genomics

5 – Managing Resilient Landscapes

•	 Is a source of studies on plant genetic variation and

gene flow 

•	 Includes many resources for identifying, collecting, developing, testing, and producing native plant materials, 

including selection guidelines and transfer zones

•	 Contains species vulnerability assessments and tools for

evaluating and predicting system resilience

•	 Is a source of management strategies that support conservation and restoration

6 – Species Endangerment

•	 Contains species vulnerability assessments and tools for

evaluating and predicting species response to disturbance

•	 Includes species distribution models for threatened and

endangered species

•	 Provides management options and strategies for species

at risk of maladaptation or extinction, such as assisted

migration or managed relocation

•	 Includes studies that use genetic approaches to conserve and restore endangered species

7 – Wildlife and Fish

•	 Includes research about climate change impacts on

wildlife and fish

•	 Contains several studies about animal translocations

and assisted migration of animal species

U.S. Forest Service Global Change Research 

Strategy, Research Elements 2009–2019

http://www.fs.fed.us/

climatechange/documents/global-change-strategy.pdf

1 – Research to enhance ecosystem 

sustainability

•	 Includes research on ecosystem health and sustainability, including assessments of climate change impacts on 

ecosystems

•	 Provides tools, applications, and technologies to improve management under a changing climate

•	 Includes genetic research that can be used to identify

plant and seed sources for restoration and conservation

•	 Provides a way to synthesize information to enhance

understanding of the changing relationships between 

climate and land disturbances

2 – Research to increase carbon sequestration

•	 Includes local, regional, and national policies and management actions for forests and wood products

•	 Provides studies that show how carbon pools may

change under different climatic scenarios

•	 Is a source of guidelines and approaches that can help

identify plant materials for sustaining and conserving 

carbon resources 

3 – Research to provide decision support

•	 Contains tools and frameworks for managing forest,

grassland, and shrubland resources in changing climates

•	 Provides information to increase our understanding of

environmental needs by species and ecosystems

•	 Highlights Forest Service expertise along with other regional and international expertise about climate change

and ecosystems

4 – Shared research needs—infrastructure, scientific collaboration, and science delivery

•	 Is a source of literature to help identify knowledge gaps,

improve data coordination and sharing, and enhance 

climate change monitoring actions

•	 Shares interdisciplinary research

•	 Serves as a central location for literature about native

plant transfer guidelines, climate change and assisted 

migration
viii USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 1

1. Anjozian LN. 2011. Checking the range for signs 

of climate change: in the past, present, and future. 

Albuquerque (NM): USDA Forest Service, Rocky Mountain 

Research Station, Grassland, Shrubland, and Desert Ecosystem 

Science Program. 8 p.

Type: Newsletter

Geographic Area: Western USA 

Compilers’ Keywords: grassland, shrubland, desert, RMRS, 

climate change, U.S. Forest Service, GSD Update

Introduction: Imagine you have access to a machine that can 

make particles move faster and faster until they approach the 

speed of light, and essentially travel through time. The machine 

might look like the Large Hadron Collider—the particle accelerator below ground in Switzerland—but instead of producing teeny, tiny, short-lived, exotic particles no one’s ever seen 

before, it transports you, a person of ample curiosity, into the 

future. You disembark your time travel machine, look around, 

and though you believe you’re in the same geography, things 

don’t look quite the same. If you began your trip somewhere in 

the interior American West where familiar grasslands, shrublands, or deserts were found, you have reason to be perplexed. 

The ecosystems of the future world have changed. What does 

that future world look like? Scientists in the Grassland, Shrubland, and Desert Ecosystems Science Program (GSD)—a unit 

of the USDA Forest Service Rocky Mountain Research Station 

(RMRS)—are not waiting for the future to arrive to have a 

look—they are working on revealing the story to us now.

Link: http://www.treesearch.fs.fed.us/pubs/39154

2. Ayling R. 2012. A history of forests and climate 

change under the IPCC. The Forestry Chronicle 88:381-382.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: climate change, forest management, 

IPCC, migration, commentary

Introduction: The Intergovernmental Panel on Climate 

Change (IPCC) was established in 1988 by two United Nations 

agencies, the World Meteorological Organization and the UN’s 

Environment Programme, to “assess the scientific, technical 

and socioeconomic information relevant for the risk of humaninduced climate change.” Its first assessment report in 1990 

was the basis for the UN Framework Convention on Climate 

Change (UNFCCC). For more than twenty years the IPCC has 

remained the most important source of scientific, technical and 

socioeconomic information for the Convention. The relationship between the UNFCCC and the IPCC is regarded by the 

international community as a “model for interaction between 

science and decision makers.”

General (1–36)—editorials, opinions, reviews, 

trends, surveys, summaries

Research (37–135)—climate change impacts, 

projections, paleoecology

Strategies (136–205)—policies, adaptation 

options, frameworks, mitigations

Resources (206–214)—tools, websites, software

Climate Change 

CHAPTER 1 • CLIMATE CHANGE • GENERAL

#. Author(s), Date. Title. Source.

Type: 

Geographic Area: 

Keywords: 

Description:

Link:

location of study

keywords written by the 

author(s) or provided by the 

compilers

ID number and

information

abstract, summary, or 

portions of the introduction 

or conclusion written by the 

author(s) or provided by the 

compilers

book, book section, conference 

paper, dissertation, government 

document, journal article, 

newsletter, presentation, report, 

thesis or web page

internet address of reference

Each reference contains the following:
2 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

5. Bierbaum R, Smith JB, Lee A, Blair M, Carter L, 

Chapin FS, Fleming P, Ruffo S, Stults M, McNeeley S, Wasley E, Verduzco L. 2012. A comprehensive 

review of climate adaptation in the United States: more 

than before, but less than needed. Mitigation and Adaptation 

Strategies for Global Change 18:361-406.

Type: Journal 

Geographic Area: USA

Keywords: adaptation process, barriers, climate change, 

mainstreaming, multiple stressors, stakeholder participation, 

successes, case studies

Abstract: We reviewed existing and planned adaptation activities of federal, tribal, state, local governments and the private sector in the United States (U.S.) to understand what types 

of adaptation activities are underway across different sectors 

and scales throughout the country. Primary sources of review 

included material officially submitted for consideration in the 

upcoming 2013 U.S. National Climate Assessment and supplemental peer-reviewed and grey literature. Although substantial 

adaptation planning is occurring in various sectors, levels of 

government and the private sector, few measures have been 

implemented and even fewer have been evaluated. Most adaptation actions to date appear to be incremental changes, not the 

transformational changes that may be needed in certain cases 

to adapt to significant changes in climate. While there appear 

to be no one-size-fits-all adaptations, there are similarities in 

approaches across scales and sectors, including mainstreaming climate considerations into existing policies and plans, and 

pursuing no- and low-regrets strategies. Despite the positive 

momentum in recent years, barriers to implementation still 

impede action in all sectors and across scales. The most significant barriers include lack of funding, policy and institutional constraints and difficulty in anticipating climate change 

given the current state of information on change. However, the 

practice of adaptation can advance through learning by doing, 

stakeholder engagements (including “listening sessions”) and 

sharing of best practices. Efforts to advance adaptation across 

the U.S. and globally will necessitate the reduction or elimination of barriers, the enhancement of information and best practice sharing mechanisms and the creation of comprehensive 

adaptation evaluation metrics.

Link: http://www.climateaccess.org/sites/default/files/Bierbaum%20et%20al_A%20comprehensive%20review%20of%20

climate%20adaptation.pdf

6. Blaser J, Gregersen H. 2013. Forests in the next 300 

years. Unasylva 64:61-73.

Type: Journal

Geographic Area: Global

Compilers’ Keywords: climate change, range shifts, forest 

ecosystems

3. Bagne KE, Finch DM. 2012. It works both ways: 

how scientists and managers join forces to conserve today’s natural resources. Albuquerque (NM): USDA Forest 

Service, Grassland, Shrubland, and Desert Ecosystem Science 

Program. 7 p.

Type: Newsletter

Geographic Area: Western USA

Compilers’ Keywords: native plant guidelines, conservation, management, GSD Update

Introduction: “Too many people fail to realize that real communication goes in both directions.” Lee Iacocca spoke these 

wise words about the failings of business, but clearly there is a 

lesson here for management of natural resources as well. Scientists are doing research every day that improves our understanding of our valued resources. But work does not stop at 

simply conducting an experiment, it needs to include effective 

communication of results that are relevant to managers. Scientists listen closely to what managers have to say and design 

studies to address questions across the diverse array of issues 

that challenge management goals today. (Features article about 

native plant transfer guidelines.)

Link: http://www.treesearch.fs.fed.us/pubs/41330

4. Bellard C, Bertelsmeier C, Leadley P, Thuiller 

W, Courchamp F. 2012. Impacts of climate change on 

the future of biodiversity. Ecology Letters 15:365-377.

Type: Journal 

Geographic Area: Global

Keywords: biodiversity, climate change, species extinctions

Abstract: Many studies in recent years have investigated the 

effects of climate change on the future of biodiversity. In this 

review, we first examine the different possible effects of climate change that can operate at individual, population, species, community, ecosystem and biome scales, notably showing that species can respond to climate change challenges by 

shifting their climatic niche along three non-exclusive axes: 

time (e.g. phenology), space (e.g. range) and self (e.g. physiology). Then, we present the principal specificities and caveats 

of the most common approaches used to estimate future biodiversity at global and sub-continental scales and we synthesize 

their results. Finally, we highlight several challenges for future 

research both in theoretical and applied realms. Overall, our 

review shows that current estimates are variable, depending on 

the method, taxonomic group, biodiversity loss metrics, spatial scales and time periods considered. Yet, the majority of 

models indicate alarming consequences for biodiversity, with 

the worst-case scenarios leading to extinction rates that would 

qualify as the sixth mass extinction in the history of the earth.

Link: http://www.ncbi.nlm.nih.gov/pubmed/22257223

CHAPTER 1 • CLIMATE CHANGE • GENERAL
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 3

support human health and well-being. The perspective serves 

as both a conservation approach and an extension of ecosystem 

management, involving the connection of ecosystem services 

to the people who benefit, in some cases with an assigned market value. We argue that the emergence of an ecosystem services perspective is timely as public interest in the state of the 

environment increases and natural resource managers face the 

reality of rapid forest ecosystem change. Forest conservation 

that considers the supply and delivery of ecosystem services 

will enhance the health and resiliency of ecosystems, engage 

and serve a broader public and attract private investment and 

leadership in a common effort to safeguard natural systems. 

Link: http://www.treesearch.fs.fed.us/pubs/31719

9. Dormann CF. 2007. Promising the future? Global 

change projections of species distributions. Basic and Applied Ecology 8:387-397.

Type: Journal 

Geographic Area: Global

Keywords: climate change, conservation management, environmental change, extrapolation, prediction, spatial statistics, 

species distribution model

Abstract: Projections of species’ distribution under global 

change (climatic and environmental) are of great scientific and 

societal relevance. They rely on a proper understanding of how 

environmental drivers determine species occurrence patterns. 

This understanding is usually derived from an analysis of the 

species’ present distribution by statistical means (species distribution models). Projections based on species distribution models make several assumptions, (such as constancy of limiting 

factors, no evolutionary adaptation to drivers, global dispersal) some of which are ecologically untenable. Also, methodological issues muddy the waters (e.g. spatial autocorrelation, 

collinearity of drivers). Here, I review the main shortcomings 

of species distribution models and species distribution projections, identify limits to their use and open a perspective on how 

to overcome some current obstacles. As a consequence, I caution biogeographers against making projections lightheartedly 

and conservation ecologists and policy makers to be aware that 

there are several unresolved problems.

Link: http://www.biom.uni-freiburg.de/mitarbeiter/dormann/publications-dormann/Dormann2007BasicApplEcol.

pdf

10. Easterling WE. 1996. Adapting North American 

agriculture to climate change in review. Agricultural and 

Forest Meteorology 80:1-53.

Type: Journal 

Geographic Area: North America

Compilers’ Keywords: climate change, agriculture, global 

warming, policy, genetic diversity, crop varieties, adaptation

Introduction (2nd paragraph): At the global scale, the 

question of whether individual trees, such as those that survive to maturity is unimportant, but the overall fate of the forests of which they are a part is crucial. Forests and trees are 

a renewable resource, providing an enormous range of goods 

and ecosystem services. In the face of expected declines in the 

availability of non-renewable resources and massive environmental change, the fate of trees and forests in the next 200–300 

years is of fundamental importance to humanity. Forests come 

and go, but in the last several hundred years there has been a 

dramatic decline. Nevertheless, there is potential to reverse this 

and to greatly increase the global forest resource. In this article, 

we consider the factors that will influence the fate of forests in 

the next 300 years, and predict a world that is more reliant than 

ever on its forests—and on its forest managers.

Link: https://www.eda.admin.ch/content/dam/deza/en/documents/Themes/resource_en_223023.pdf

7. Brown JH, Gibson AC. 1983. Biogeography. St. 

Louis (MO): C. V. Mosby. 643 p. 

Type: Book

Geographic Area: Global

Compilers’ Keywords: biodiversity, evolutionary processes, 

geography

Compilers’ Summary: Biogeography (2nd edition) combines ecological and historical perspectives to show how contemporary environments, Earth history and evolutionary processes have shaped the distributions of species and the patterns 

of biodiversity. It illustrates general patterns and processes 

using examples from different groups of plants and animals 

from diverse habitats and geographic regions. Written primarily for use in undergraduate and graduate courses in plant and/

or animal geography, the book serves as a general synthesis 

and reference as well.

8. Collins S, Larry E. 2007. Caring for our natural 

assets: an ecosystem services perspective. Portland (OR): 

USDA Forest Service, Pacific Northwest Region. General 

Technical Report PNW-GTR-733. 11 p.

Type: Government Document

Geographic Area: USA

Keywords: ecosystem services, ecosystem management, natural capital, climate change, human well-being

Abstract: Global attention to climate change has advanced an 

awareness of human impacts on the environment. Progressing 

more slowly is recognition of the critical link between forest 

ecosystems and human welfare. Forests provide a number of 

societal benefits or ecosystem services, such as water purification, climate and flood regulation, recreational opportunities 

and spiritual fulfillment. This paper examines an emerging 

perspective that describes ecosystems as natural assets that 

CHAPTER 1 • CLIMATE CHANGE • GENERAL
4 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Type: Government Document

Geographic Area: Global

Compilers’ Keywords: climate change, ecosystems, biodiversity, ecosystem services, impacts, management, assessment

Introduction: Climate change affects the living world, including people, through changes in ecosystems, biodiversity 

and ecosystem services. Ecosystems entail all the living things 

in a particular area, as well as the non-living things with which 

they interact, such as air, soil, water and sunlight. Biodiversity 

refers to the variety of life, including the number of species, 

life forms, genetic types and habitats and biomes (which are 

characteristic groupings of plant and animal species found in 

a particular climate). Biodiversity and ecosystems produce a 

rich array of benefits that people depend on, including fisheries, drinking water, fertile soils for growing crops, climate 

regulation, inspiration and aesthetic and cultural values. These 

benefits are called “ecosystem services”—some of which, 

like food and fisheries, are more easily quantified than others, such as climate regulation or cultural values. Ecosystem 

services translate into jobs, economic growth, health and human well-being. Although ecosystems and ecosystem services 

are what we interact with every day, their linkage to climate 

change can be elusive because they are influenced by so many 

additional entangled factors. Ecosystem perturbations driven 

by climate change have direct human impacts, including reduced water supply and quality, the loss of iconic species and 

landscapes, distorted rhythms of nature and the potential for 

extreme events to overcome the regulating services of ecosystems. Even with these well-documented ecosystem impacts, 

it is often difficult to quantify human vulnerability that results 

from shifts in ecosystem processes and services. For example, 

although it is straightforward to predict how precipitation will 

change water flow; it is much harder to pinpoint which farms 

and cities will be at risk of running out of water, and even more 

difficult to say how people will be affected by the loss of a favorite fishing spot or a wildflower that no longer blooms in the 

spring. A better understanding of how everything from altered 

water flows to the loss of wildflowers matters to people may be 

key to managing ecosystems in a way that promotes resilience 

to climate change.

Link: http://ncadac.globalchange.gov/download/NCAJan11-

2013-publicreviewdraft-chap8-ecosystems.pdf

12. Hajjar R, McGuigan E, Moshofsky M, Kozak 

RA. 2014. Opinions on strategies for forest adaptation to future climate conditions in western Canada: surveys of the general 

public and leaders of forest-dependent communities. Canadian 

Journal of Forest Research 44:1525-1533.

Type: Journal

Geographic Area: Western Canada

Keywords: assisted migration, biotechnology, public opinion, 

reforestation, climate change adaptation

Abstract: The adaptability of North American agriculture to 

climate change is assessed through a review of current literature. A baseline of North American agriculture without climate 

change suggests that farming faces serious challenges in the 

future (e.g. declining domestic demand, loss of comparative 

advantage, rising environmental costs). Climate change adjustments at the farm-level and in government policy, including 

international trade policy, are inventoried from the literature. 

The adaptive potential of agriculture is demonstrated historically with situations that are analogous to climate change, including the translocation of crops across natural climate gradients, the rapid introduction of new crops such as soybeans 

in the U.S. and canola in Canada and resource substitutions 

prompted by changes in prices of production inputs. A wide 

selection of modeling studies is reviewed, which in net, suggests several agronomic and economic adaptation strategies 

that are available to agriculture. Agronomic strategies include 

changes in crop varieties and species, timing of operations and 

land management including irrigation. Economic strategies 

include investment in new technologies, infrastructure and labor and shifts in international trade. Overall, such agronomic 

strategies were found to offset either partially or completely 

the loss of productivity caused by climate change. Economic 

adaptations were found to render the agricultural costs of climate change small by comparison with the overall expansion 

of agricultural production. New avenues of adaptive research 

are recommended including the formalization of the incorporation of adaptation strategies into modeling, linkage of adaptation to the terrestrial carbon cycle, anticipation of future 

technologies, attention to scaling from in situ modeling to the 

landscape scale, expansion of data sets and the measurement 

and modeling of unpriced costs. The final assessment is that 

climate change should not pose an insurmountable obstacle to 

North American agriculture. The portfolio of assets needed to 

adapt is large in terms of land, water, energy, genetic diversity, 

physical infrastructure and human resources, research capacity 

and information systems and political institutions and world 

trade—the research reviewed here gives ample evidence of the 

ability of agriculture to utilize such assets. In conclusion, the 

apparent efficiency with which North American agriculture 

may adapt to climate changes provides little inducement for 

diverting agricultural adaptation resources to efforts to slow or 

halt climate changes.

Link: http://www.climateknowledge.org/figures/Rood_Climate_Change_AOSS480_Documents/Easterling_North_

American_Agriculture_AgricForMet_1996.pdf

11. Groffman PM, Kareiva P, Carter S, Grimm NB, 

Lawler JJ, Mack M, Matzek V, Tallis H. 2013. Ecosystems, biodiversity, and ecosystem services (Draft for public comment). Chapter 8. In: NCADAC Draft Climate Assessment Report. Washington (DC): U.S. Global Change Research 

Program. p 291-330. 

CHAPTER 1 • CLIMATE CHANGE • GENERAL
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 5

14. Hodgkin T, Bordoni P. 2012. Climate change and 

the conservation of plant genetic resources. Journal of Crop 

Improvement 26:329-345.

Type: Journal 

Geographic Area: Global

Keywords: plant genetic resources, conservation, ex situ, in 

situ, adaptation

Abstract: Over the next decades, agricultural production 

practices will change significantly and become more sustainable while they also respond to the need to contribute to reducing malnutrition and hunger and meeting the challenges of 

climate change. The enhanced use of agricultural biodiversity 

will play an essential role in this process, providing improved 

adaptability and resiliencies in agro-ecosystems. Plant genetic 

resources, a major component of agricultural biodiversity, play 

a key role in improving agricultural production and productivity. They are also essential to coping with climate change. As 

a result of climate change, increased efforts will be needed to 

conserve the diversity of crops and their wild relatives, and 

both in situ and ex situ conservation strategies will have to be 

adapted to meet changing environmental conditions and the 

need to secure biodiversity threatened by changing climate 

and altered production practices. Improved use of plant genetic resources will be essential, and this is likely to require 

increased national and international movements of resources 

to ensure that adapted germplasm is available to meet changing production environments. Greater emphasis will also need 

to be placed on evaluation for resistance to biotic and abiotic 

stresses and on properties such as adaptability, plasticity and 

resilience, which can help maintain productivity under changing environmental conditions. 

15. Hopkins AD. 1938. Bioclimatics: a science of life and 

climate relations. Washington (DC): U.S. Government Printing 

Office. Miscellaneous Publication No. 280. 199 p.

Type: Government Document

Geographic Area: Global

Compilers’ Keywords: climatic adaptation, Hopkin’s law, 

seasonal variation, species distribution

Introduction: Bioclimatics is a science of relations between 

life, climate, seasons and geographic distribution. It deals with 

fundamental laws, principles, systems and methods of application in general research and economic practice, and has 

special reference to the major and minor effects of the major 

astronomic and terrestrial laws of causation, as represented by 

the variable phenomena of life, climate and seasons, relative to 

the geographic coordinates as expressed, measured and interpreted in units of time, temperature, and distance. The science 

differs from the other branches of biology, climatology, and 

geography in that it is based on the bioclimatic (and related) 

laws and principles, and in that it deals more specifically with 

Abstract: Two province-wide surveys of residents in Alberta 

and British Columbia were conducted to assess the acceptability of a range of reforestation strategies—many of which 

revolve around biotechnology—that could be used to aid western Canada’s forests in adapting to future climate change. The 

opinions of leaders of forest-dependent communities were also 

sought to evaluate how well they align with those of the public 

at large. Results show that the views of the general public and 

community leaders correspond. There is a low acceptance for 

a “do nothing” strategy that allows climate change to run its 

course without any human intervention; high acceptance of replanting with local seeds; a decreasing acceptance of strategies 

that involve more manipulation such as breeding, using nonlocal seeds and moving seeds outside of a species’ natural range; 

and a low acceptance of genetically engineered solutions. However, a high proportion of respondents changed their answers 

when told that a particular strategy would lead to either favourable or unfavourable outcomes related to socioeconomics of 

forest-dependent communities, forest aesthetics, pest, disease 

and fire outbreaks. We conclude that a meaningful and participatory dialogue on forest adaptation strategies in the face of climate change can only emerge if residents and other interested 

stakeholders have an adequate understanding of current forest 

management practices, proposed reforestation strategies, the 

role of technological interventions and the values and services 

for which western Canada’s forests are to be managed.

Link: http://www.nrcresearchpress.com/doi/abs/10.1139/cjfr2014-0142#.VTiDQZgcRLM

13. Harris JA, Hobbs RJ, Higgs E, Aronson J. 2006.

Ecological restoration and global climate change. Restoration 

Ecology 14:170-176.

Type: Journal 

Geographic Area: Global

Keywords: climate change, ecosystem change, ecosystem 

function, historical ecosystem, restoration goals

Abstract: There is an increasing consensus that global climate 

change occurs and that potential changes in climate are likely 

to have important regional consequences for biota and ecosystems. Ecological restoration, including (re)-afforestation 

and rehabilitation of degraded land, is included in the array 

of potential human responses to climate change. However, the 

implications of climate change for the broader practice of ecological restoration must be considered. In particular, the usefulness of historical ecosystem conditions as targets and references must be set against the likelihood that restoring these 

historic ecosystems is unlikely to be easy, or even possible, in 

the changed biophysical conditions of the future. We suggest 

that more consideration and debate needs to be directed at the 

implications of climate change for restoration practice.

Link: http://nctc.fws.gov/CSP/Resources/fwca/Climate%20

Change/Ecological%20restoration%20and%20CC.pdf

CHAPTER 1 • CLIMATE CHANGE • GENERAL
6 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

benefits. Section C considers principles for effective adaptation and the broader interactions among adaptation, mitigation, 

and sustainable development.

Link: https://ipcc-wg2.gov/AR5/images/uploads/WG2AR5_

SPM_FINAL.pdf

17. Jansen E, Overpeck J, Briffa KR, Duplessy J-C, 

Joos F, Masson-Delmotte V, Olago D, Otto-Bliesner 

B, Peltier WR, Rahmstorf S, Ramesh R, Raynaud 

D, Rind D, Solomina O, Villalba R, Zhang D. 2007.

Palaeoclimate. Chapter 6. In: Solomon S, Qin D, Manning M, 

Chen Z, Marquis M, Averyt KB, Tignor M, Miller HL, editors. 

Climate Change 2007: The Physical Science Basis. Cambridge, 

United Kingdom: Cambridge University Press. p 433-498. 

Type: Book Section

Geographic Area: Global

Compilers’ Keywords: climate change, migration, emissions scenarios, paleoecology, palaeoclimate, climate models, 

ice age, warming, greenhouse gases

Introduction: This chapter assesses palaeoclimatic data and 

knowledge of how the climate system changes over interannual to millennial time scales, and how well these variations can 

be simulated with climate models. Additional palaeoclimatic 

perspectives are included in other chapters. Palaeoclimate science has made significant advances since the 1970s, when a 

primary focus was on the origin of the ice ages, the possibility 

of an imminent future ice age, and the first explorations of the 

so-called Little Ice Age and Medieval Warm Period. Even in 

the first IPCC assessment (IPCC, 1990), many climatic variations prior to the instrumental record were not that well known 

or understood. Fifteen years later, understanding is much improved, more quantitative and better integrated with respect to 

observations and modelling. After a brief overview of palaeoclimatic methods, including their strengths and weaknesses, 

this chapter examines the palaeoclimatic record in chronological order, from oldest to youngest. This approach was selected 

because the climate system varies and changes over all time 

scales, and it is instructive to understand the contributions that 

lower-frequency patterns of climate change might make in influencing higher-frequency patterns of variability and change. 

In addition, an examination of how the climate system has responded to large changes in climate forcing in the past is useful 

in assessing how the same climate system might respond to the 

large anticipated forcing changes in the future. 

18. Joyce LA, Running SW, Breshears DD, Dale 

VH, Malmsheimer RW, Sampson RN, Sohngen B, 

Woodall CW. 2013. Forestry. Chapter 7. In: NCADAC 

Draft Climate Assessment Report. Washington (DC): U.S. 

Global Change Research Program. p 263-289. 

Type: Government Document

Geographic Area: Global

fundamentals and methods of application relative to the phenomena of seasons, climates, and geographic distribution. It is 

not, therefore, a branch of any one of the major or minor sciences, but is intended to be supplementary to all in contributing 

information on fundamentals, and to serve a specific purpose. 

Link:http://books.google.com/books?hl=en&lr

&id=OpE9wLvCq8oC&oi=fnd&pg=PA1&dq=B

ioclimatics:+A+science+of+life+and+climate+re

lations&ots=xoklNA0odR&sig=kwq_AHk28qk7pDdQDLinbQtzYA#v=onepage&q=Bioclimatics%3A%20

A%20science%20of%20life%20and%20climate%20

relations&f=false

16. Intergovernmental Panel on Climate 

Change [IPCC]. 2014. Summary for policymakers.

In: Field CB, Barros VR, Dokken DJ, Mach KJ, Mastrandrea 

MD, Bilir TE, Chatterjee M, Ebi KL, Estrada YO, Genova RC, 

Girma B, Kissel ES, Levy AN, MacCracken S, Mastrandrea 

PR, White LL, editors. Climate Change 2014: Impacts, 

Adaptation, and Vulnerability. Part A: Global and Sectoral 

Aspects. Contribution of working group II to the fifth assessment report of the Intergovernmental Panel on Climate 

Change. New York (NY): Cambridge University Press. 32 p. 

Type: Government Document

Geographic Area: Global

Compilers’ Keywords: climate change, natural resources, 

human impacts, socioeconomic processes, land-use change, 

risks, vulnerability, exposure, hazards, decisionmaking

Introduction: Human interference with the climate system is 

occurring, and climate change poses risks for human and natural systems. The assessment of impacts, adaptation, and vulnerability in the Working Group II contribution to the IPCC’s 

Fifth Assessment Report (WGII AR5) evaluates how patterns of risks and potential benefits are shifting due to climate 

change. It considers how impacts and risks related to climate 

change can be reduced and managed through adaptation and 

mitigation. The report assesses needs, options, opportunities, 

constraints, resilience, limits, and other aspects associated with 

adaptation. Climate change involves complex interactions and 

changing likelihoods of diverse impacts. A focus on risk, which 

is new in this report, supports decision making in the context of 

climate change and complements other elements of the report. 

People and societies may perceive or rank risks and potential 

benefits differently, given diverse values and goals. Compared 

to past WGII reports, the WGII AR5 assesses a substantially 

larger knowledge base of relevant scientific, technical, and 

socioeconomic literature. Increased literature has facilitated 

comprehensive assessment across a broader set of topics and 

sectors, with expanded coverage of human systems, adaptation, and the ocean. Section A of this summary characterizes 

observed impacts, vulnerability and exposure, and adaptive responses to date. Section B examines future risks and potential 

CHAPTER 1 • CLIMATE CHANGE • GENERAL
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 7

around sources of organisms; 55% prefer native source material 

from close sources and 32% prefer them from similar habitats, 

not necessarily the closest source. A majority prefer organisms 

from a mix of populations (71%) compared to single sources 

(10%). Regarding genetic issues during captive production, 

practitioners ranked loss of genetic diversity above domestication selection or outbreeding depression. A strong majority 

(89%) of respondents were convinced that climate change is 

underway, and many (75%) believed that the practice of restoration should be modified to anticipate it. For example, the 

practice of setting restoration goals to match historic conditions 

may become infrequent or no longer used by many (65%) if 

climate changes. Many practitioners were cautious about moving species in response to climate change. For example, moving 

species beyond their historic range was unpopular (10% said 

‘yes’ while 30% chose ‘maybe’) unless the species faces extinction. In contrast, moving species within their historic range but 

between watersheds was acceptable to many (56% said ‘yes’ 

and 22% said ‘maybe’). Also, respondents were somewhat 

(42%) or very (55%) concerned that species moved in response 

to climate change could become invasive in a new area. These 

survey results suggest many commonalities among restorationists regarding the major issues they face while conducting restoration, obtaining organisms, and responding to climate change. 

In particular, there is widespread agreement that restoration 

should respond to shifting climates but exactly how to respond 

remains controversial.

Link: http://www.nativeseed.info/2010/presentations/

Kaye%20-%20restoration%20and%20climate%20change.pdf

20. Kujala H, Burgman MA, Moilanen A. 2012.

Treatment of uncertainty in conservation under climate change. 

Conservation Letters 6:73-85.

Type: Journal 

Geographic Area: Global

Keywords: biodiversity, conservation planning, epistemic 

uncertainty, global warming, human decision uncertainty, linguistic uncertainty, uncertainty analysis

Abstract: Climate change is an important threat to biodiversity 

globally, but there are major uncertainties associated with its 

magnitude and ecological consequences. Here, we investigate 

how three major classes of uncertainty, linguistic uncertainty, 

epistemic uncertainty (uncertainty about facts), and human 

decision uncertainty, have been accounted for in scientific literature about climate change. Some sources of uncertainty are 

poorly characterized and epistemic uncertainty is much more 

commonly treated than linguistic or human decision uncertainty. Furthermore, we show that linguistic and human decision uncertainties are relatively better treated in the literature on 

sociopolitics or economics than in natural sciences, which often 

overlook communication between stakeholders and socioeconomic consequences. As uncertainty can significantly influence 

Compilers’ Keywords: forest management, climate change, 

impacts, ecosystem services

Key Messages: Climate change is increasing the vulnerability 

of forests to ecosystem change and tree mortality through fire, 

insect infestations, drought, and disease outbreaks. Western 

U.S. forests are particularly vulnerable to increased wildfire 

and insect outbreaks; eastern forests have smaller disturbances 

but could be more sensitive to periodic drought. U.S. forests 

currently absorb about 13% of all carbon dioxide (CO2) emitted by fossil fuel burning in the U.S. Climate change, combined 

with current societal trends regarding land use and forest management, is projected to reduce forest CO2 uptake. Bioenergy 

is an emerging new market for wood; with higher wood prices, 

development of a market in salvaged wood from trees killed 

by drought, insects, and fire could help finance salvage and 

restoration activities and reduce U.S. fossil fuel consumption. 

However, the environmental and socioeconomic consequences 

of bioenergy production vary greatly with region and intensity 

of human management. The changing nature of private forestland ownership, globalization of forestry markets, emerging markets for bioenergy, and U.S. climate change policy will 

all influence forest management responses to climate change. 

However, development of and better access to practical and 

timely information for managers to consider in choosing adaptation and mitigation options will facilitate management of 

public and private forestland.

Link: http://ncadac.globalchange.gov/download/NCAJan11-

2013-publicreviewdraft-chap7-forestry.pdf

19. Kaye TN. 2010. What do we think? An international 

survey on restoration and climate change. National Native Seed 

Conference: Native Plant Materials Development, Production, 

and Use for Habitat Restoration. Corvallis (OR): Institute for 

Applied Ecology. 37 p.

Type: Presentation

Geographic Area: USA

Compilers’ Keywords: climate change, survey, restoration, 

paradigm, managers, opinion

Abstract: Climate change may be the defining challenge to the 

field of restoration ecology this century. How does the wider restoration community currently approach the challenges of habitat and species restoration, and how is this approach likely to 

shift if the climate changes locally and globally? Understanding 

how people conduct or support restoration is crucial to engaging in discussions that move our field forward in the face of 

changing environments. We conducted an international survey 

of over 1000 restoration practitioners and ecologists to gather 

information on their perspectives about these issues. We asked 

a series of twenty questions focused on the restoration process 

and obtaining organisms for restoration, climate change, and 

moving species in response to climate change. Survey results 

suggest that practitioners are somewhat split in their preferences 

CHAPTER 1 • CLIMATE CHANGE • GENERAL
8 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

on forest ecosystems. The obvious relationships—like severe 

drought and forest fires, or warmer decades and increased survival of seedlings at upper tree line—have been well studied. 

Other effects, such as the combined role of climate in insect 

life cycles and their tree hosts’ vulnerability, are just beginning to be understood. So the idea that climate variability plays 

a role in forests is not new, nor is the idea that understanding that variability is critical for successful planning in forest 

management and forestry. But events like the 2002 Biscuit fire 

in Oregon, nearly 500,000 acres, or mountain pine beetle epidemics that impact entire watersheds synchronously over the 

West, give us pause.

Link: http://www.forestry.org/media/docs/westernforester/2009/jan09.pdf

23. Næss MW. 2013. Climate change, risk management 

and the end of Nomadic pastoralism. International Journal of 

Sustainable Development & World Ecology 20:123-133.

Type: Journal 

Geographic Area: Tibet, Asia

Keywords: fragmentation, climate change, density dependence, risk, nomadic pastoralism, grazing patterns, Chang 

Tang, Tibetan Plateau, China

Abstract: Mobility has been argued to be the single factor 

explaining why some pastoralists do relatively well during extreme climatic events, while others do not, because mobility 

works by taking advantage of the spatial and temporal structure of resource failure by moving away from scarcity towards 

abundance. In spite of this, a common governmental management strategy is to resettle pastoral populations and thereby 

significantly reduce mobility. By revealing the underlying 

logic of mobility for Tibetan pastoralists, this paper questions 

official policy that aims at privatizing communally owned 

rangelands since it reduces pastoral flexibility and access to 

key resources. This is especially pertinent in the face of climate 

change. While little is known as to the specifics of how climate 

change will affect nomadic pastoralists, environmental variability is likely to increase. Consequently, policies resulting 

in decreased mobility may exacerbate the negative effects of 

climate change because of a positive feedback between climate 

and negative density dependence.

24. Nakicenovic N, Davidson O, Davis G, Grubler 

A, Kram T, Lebre La Rovere E, Metz B, Morita 

T, Pepper W, Pitcher H, Sankovski A, Shukla P, 

Swart R, Watson R, Zhou D. 2000. Summary for policy 

makers: emissions scenarios, a special report of Working Group 

III of the Intergovernmental Panel on Climate Change. PNNLSA-39650. 27 p.

Type: Government Document

Geographic Area: Global

implementation of conservation, we discuss uncertainties associated with some commonly proposed conservation adaptation actions to mitigate climate change. There may be major 

differences between strategies, with implications on how they 

should be viewed in conservation planning. We conclude that 

evaluating conservation strategies in terms of different types of 

uncertainty will facilitate communication between disciplines 

and stakeholders. While accounting for uncertainties in a quantitative manner is difficult and data demanding, even qualitative 

appreciation about the uncertainties inherent in conservation 

strategies can facilitate and improve decision making. 

Link: https://tuhat.halvi.helsinki.fi/portal/files/26044830/

Kujala_et_al_2013_CL.pdf

21. Lenart M, Jones C. 2014. Perceptions on climate 

change correlate with willingness to undertake some forestry adaptation and mitigation practices. Journal of Forestry 

112:553-563.

Type: Journal

Geographic Area: USA

Keywords: climate change, adaptation, mitigation, climate 

records, forest management

Abstract: Results from a survey taken anonymously by 1,029 

U.S. -based respondents in forest management and academia 

highlight how individuals’ perceptions about climate change 

correlated with their willingness to consider management 

prescriptions for forest ecosystems. About two-thirds of respondents identified themselves as having a professional land 

management role, and most of the remainder specialized in 

research or education. In most cases, respondents’ willingness 

to try specific adaptation or mitigation measures related to the 

degree to which they agreed that “climate change is occurring 

because of human activities that release greenhouse gases to 

the atmosphere.” The survey results are considered with a focus on which proposed practices and types of climate information received the most acceptance or resistance. For instance, 

respondents across the spectrum of climate change perceptions 

supported efforts to thin overly dense forests and opposed options to sequester carbon by promoting the woody invasion of 

grasslands or ignoring biodiversity.

Link: http://www.ingentaconnect.com/content/saf/

jof/2014/00000112/00000006/art00003

22. Littell J, Peterson D. 2009. Pacific Northwest forests 

and climate. Western Forester 54:1-24.

Type: Newsletter

Geographic Area: USA

Compilers’ Keywords: climate change, forest ecosystems, 

genotypes, forest management, adaptation, diversity

Introduction: We already know from over a century of observations in Western forests that climate exerts strong influences 

CHAPTER 1 • CLIMATE CHANGE • GENERAL
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 9

research that would inform adaptation of BC’s current forest 

management policies to a changing climate. A key goal of the 

council was to maximize the utility of the research to inform 

provincial policy. The eight- step process that we developed to 

achieve this goal is described in this paper. In roughly chronological order, the eight steps were: determining the research 

needed to inform policy, connecting scientists and policy-makers, requiring interdisciplinary teams including both natural 

and social scientists and relevant stakeholders, assessing proposals for their value to inform policy, fostering scientific excellence, fostering ongoing communication between scientists 

and policy-makers, tailoring communication to policy-makers, 

and disseminating the policy relevant outcomes in a timely and 

targeted manner. Based on the FFESC experience, we suggest 

best practices for engaging policy-makers in research and scientists in policy development and adaptation.

Link: http://pubs.cif-ifc.org/doi/abs/10.5558/tfc2014-014

27. Regniere J, Bentz B. 2008. Mountain pine beetle 

and climate change. In: McManus K, Gottschalk KW, editors. 

Proceedings 19th USDA Interagency Research Forum on 

Invasive Species 2008. Newtown Square (PA): USDA Forest 

Service, Northern Research Station. General Technical Report 

NRS-P-36. p 63-64.

Type: Government Document

Geographic Area: Western North America

Compilers’ Keywords: climate change, mountain pine beetle, management, forestry, invasive species, boreal forest

Abstract: The mountain pine beetle (MPB) (Dendroctonus 

ponderosae) is a native insect of pine forests in western North 

America. While it has a broad geographical distribution, it has 

been historically confined to the western side of the continent, 

in the U.S. by the distribution of its pine hosts and in the northern half of British Columbia in western Canada by the geoclimatic barrier of the Rockies. Since the early to mid-1990s, an 

outbreak of MPB has reached unprecedented levels in terms 

of acreages and numbers of pine trees, in particular lodgepole 

pine, killed throughout its range, most notably in Colorado and 

British Columbia. The MPB is also causing very high mortality among whitebark and limber pines at high elevations. 

Historical records from the past 100 years suggest these ecosystems have had pulses of MPB-caused mortality but not at 

levels currently being observed. Since 2006, MPB has extended its range into the Peace River area of north-central Alberta. 

Climate change may well be involved in this recent northeastward and upward range expansion. There is ample and mounting evidence of similar latitudinal and altitudinal shifts in 

insect distributions throughout the world, many convincingly 

Linked to climate change. The main concern at this time is the 

likelihood that this insect will continue spreading east into the 

pines of Canada’s boreal forest, eventually reaching the eastern 

provinces and threatening the pines growing on the Atlantic 

Compilers’ Keywords: climate change, emissions scenarios, 

policy, IPCC

Preface: The Intergovernmental Panel on Climate Change 

(IPCC) was established jointly by the World Meteorological 

Organization (WMO) and the United Nations Environment 

Programme (UNEP) to assess periodically the science, impacts 

and socioeconomics of climate change and of adaptation and 

mitigation options. The IPCC provides, on request, scientific 

and technical advice to the Conference of Parties (COP) to the 

United Nations Framework Convention on Climate Change 

(UNFCCC) and its bodies. In response to a 1994 evaluation of 

the earlier IPCC IS92 emissions scenarios, the 1996 Plenary of 

the IPCC requested this Special Report on Emissions Scenarios 

(SRES) (see Appendix I for the Terms of Reference). This report was accepted by the Working Group III (WGIII) plenary 

session in March 2000. The long-term nature and uncertainty 

of climate change and its driving forces require scenarios that 

extend to the end of the 21st century. This Report describes the 

new scenarios and how they were developed.

Link: https://docs.google.com/a/mtu.edu/file/

d/0B1gFp6Ioo3akcXQxMnNsWnRlSTA/edit?usp=drive_

web&urp=http://www.ipcc.ch/publications_and_data/

publicati&pli=1

25. Prescott C. 2012. Forests, climate change, and science: 

we need to do better. The Forestry Chronicle 88:371-372.

Type: Journal 

Geographic Area: Western Canada

Compilers’ Keywords: editorial, climate change, forest, 

Canada

Introduction: To Canadian foresters, climate change is not 

a vague concept of something that might happen in the future. 

Canadian foresters are planting and managing forests that will 

be harvested or otherwise enjoyed decades in the future, so 

considering possible future climates is part of their day-to day 

work. It is therefore essential that Canadian foresters be provided with the best possible advice to help them plan and manage forests that are resilient in uncertain future climates. 

26. Prescott C, Weese K. 2014. Crossing the divide: 

engaging scientists and policy-makers in adapting forest management to climate change in British Columbia. The Forestry 

Chronicle 90:89-95.

Type: Journal 

Geographic Area: Western Canada

Keywords: science, policy development, forest management, 

climate change, interdisciplinary research, client-based research, best practices

Abstract: The Future Forest Ecosystems Scientific Council 

(FFESC) was created in 2008 following a one-time allocation of funding from the BC provincial government to support 

CHAPTER 1 • CLIMATE CHANGE • GENERAL
10 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

resource management adaptation and preparedness in a period 

of accelerating environmental change.

Link: http://www.treesearch.fs.fed.us/pubs/46127

30. Sutherland WJ, Clout M, Cote IM, Daszak P, 

Depledge MH, Fellman L, Fleishman E, Garthwaite 

R, Gibbons DW, De Lurio J, Impey AJ, Lickorish 

F, Lindenmayer D, Madgwick J, Margerison C, 

Maynard T, Peck LS, Pretty J, Prior S, Redford 

KH, Scharlemann JP, Spalding M, Watkinson AR. 

2010. A horizon scan of global conservation issues for 2010.

Trends in Ecology and Evolution 25:1-7.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: aerosols, animals, atmosphere, charcoal, climate change, conservation, ecosystem, environmental 

pollution, forecasting, genetic engineering, oceans and seas, 

nitrogen, oxygen, seawater, telemetry, volcanic eruptions

Abstract: Horizon scanning identifies emerging issues in a 

given field sufficiently early to conduct research to inform policy and practice. Our group of horizon scanners, including academics and researchers, convened to identify fifteen nascent 

issues that could affect the conservation of biological diversity. 

These include the impacts of and potential human responses 

to climate change, novel biological and digital technologies, 

novel pollutants and invasive species. We expect to repeat this 

process and collation annually.

Link: http://www.ncbi.nlm.nih.gov/pubmed/19939492

31. Swift K. 2010. Searching for sustainability in forest 

management: is good silviculture the key? Journal of Ecosystems 

and Management 12:7-9.

Type: Journal 

Geographic Area: Western North America

Compilers’ Keywords: climate change, forest management, 

Canada, United States, workshop

Introduction: I am pleased to summarize the first session 

from the 2010 Winter Southern Interior Silviculture Committee 

(SISCO) Workshop. As most are aware, the financial situation that a lot of us find ourselves in has affected our ability 

to attend many face-to-face events; however, although numbers were down, SISCO was able to deliver an informative 

one-and-a-half-day session that focused on climate change, 

sustainable forest management, and issues and concerns of 

small firms and operators. This article provides an update 

on the workshop’s opening plenary session, titled “Climate 

Change—The Underlying Reality that Shapes Forest Planning 

and Management,” and the latest science that can help inform 

decisions.

side of the continent all the way into the Southern U.S. Because 

of this recent incursion at the gates of the Canadian boreal forest, MPB is being viewed as a potential invading species in 

eastern pine ecosystems. It could be viewed as an invader into 

the high-elevation whitebark pine ecosystems as well.

Link: http://www.treesearch.fs.fed.us/pubs/19347

28. Richardson DM. 2012. Conservation biogeography: what’s hot and what’s not? Diversity and Distributions 

18:319-322.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: conservation, diversity, editorial, climate change, invasion biology, biogeography

Conclusion: Conservation biogeography is now well established as an exciting and important field in the cloud of 

subdisciplines associated with biogeography, ecology and environmental management. The challenges in biodiversity conservation are increasing rapidly, and innovative approaches and 

tools are urgently needed. I hope you will agree that Diversity 

and Distributions has carved an important niche for itself in the 

growing number of journals dealing with conservation.

29. Sample VA, Bixler RP, editors. 2013. Forest conservation and management in the Anthropocene: adaptation of 

science, policy, and practices. Fort Collins (CO): USDA Forest 

Service, Rocky Mountain Research Station. Proceedings 

RMRS-P-71. 494 p. 

Type: Government Document

Geographic Area: Global

Compilers’ Keywords: forest management, adaptation, 

climate change, conservation, restoration, forest ecosystems, 

Pinchot

Abstract: Climate change is but one aspect of the 

Anthropocene, a new epoch in which the effects of human activities have become the predominant force in the global biosphere. More than just an overlay on the traditional concerns 

of sustainable natural resource management, the uncertainties 

associated with these effects are creating a “no-analog future” 

in which much of the existing science relating to the functioning and response of forest ecosystems—which serves as the 

fundamental basis for current forest management practices 

and policies—must be reconsidered. In these collected papers, 

leading scientists, resource managers and policy specialists 

explore the implications of climate change and other manifestations of the Anthropocene on the management of wildlife 

habitat, biodiversity, water, and other resources, with particular 

attention to the effects of wildfire. Recommendations include 

the need for a supporting institutional, legal, and policy framework that is not just different but more dynamic, to facilitate 

CHAPTER 1 • CLIMATE CHANGE • GENERAL
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 11

ecosystems, especially the accelerating loss of natural biological communities, and the alteration of the chemical composition of the atmosphere. 

34. Walsh J, Wuebbles D, Hayhoe K, Kunkel K, 

Somerville R, Stephens G. 2013. Our changing climate. 

Chapter 2. In: NCADAC Draft Climate Assessment Report. 

Washington (DC): U.S. Global Change Research Program. p 

25-103. 

Type: Government Document

Geographic Area: Global

Compilers’ Keywords: climate change, anthropogenic disturbance, precipitation, temperature, frost-free season, hurricanes, heat wave

Introduction: This chapter summarizes how climate is changing, why it is changing, and what is projected for the future. 

While the focus is on changes in the United States, the need 

to provide context requires a broader geographical perspective 

in some parts of the discussion. Additional geographic detail 

is presented in the regional chapters of this report. Since the 

previous national climate assessment was published in 2009, 

the climate has continued to change, with resulting effects on 

the U.S. The trends described in the 2009 report have continued, and our understanding of the data and ability to model the 

many facets of the climate system have increased substantially. 

Several noteworthy advances are mentioned in the chapter.

Link: http://ncadac.globalchange.gov/download/NCAJan11-

2013-publicreviewdraft-chap2-climate.pdf

35. Welch D. 2005. What should protected areas managers do in the face of climate change? The George Wright Forum 

22:75-93.

Type: Journal 

Geographic Area: Canada

Compilers’ Keywords: climate change, parks, adaptive 

management, public concern

Compilers’ Summary: This paper describes the impact of 

past, current and future climate change and how managers 

should approach protecting land and adaptive management to 

promote sustainable and resilient habitats. The author uses examples from Parks Canada to illustrate why and when to adapt, 

how to adapt, what to do (i.e., goals and actions) and what not 

to do. Adapting to climate change and protecting natural areas 

required active involvement and collaboration among scientists, managers, the public and stakeholders.

Link: http://www.climateaccess.org/sites/default/files/Welch_

What%20Should%20Protected%20Areas%20Managers%20

Do.pdf

32. U.S. EPA. 2012. Climate change indicators in the 

United States. Washington (DC): United States Environmental 

Protection Agency. EPA 430-F-12-032. 4 p.

Type: Government Document

Geographic Area: USA

Compilers’ Keywords: indicators, global trends

Compilers’ Summary: The Earth’s climate is changing. 

Temperatures are rising, snow and rainfall patterns are shifting, and more extreme climate events—like heavy rainstorms and record high temperatures—are already affecting 

society and ecosystems. Scientists are confident that many 

of the observed changes in the climate can be Linked to 

the increase in greenhouse gases in the atmosphere, caused 

largely by people burning fossil fuels to generate electricity, heat and cool buildings, and power vehicles. The EPA is 

working with many other organizations to better understand 

the causes and effects of climate change. With help from 

these partners, the EPA has compiled a set of 26 indicators 

tracking signs of climate change. Most of these indicators 

focus on the United States, but some include global trends 

to provide context or a basis for comparison. These indicators represent a selected set of key climate change measurements, and are not an exhaustive group of all climate change 

indicators. The EPA’s indicators are based on peer-reviewed 

data from various government agencies, academic institutions, and other organizations. EPA selected these indicators 

based on the quality of the data and other criteria.

Link: http://www.epa.gov/climatechange/science/indicators/

33. Walker BH, Steffen WL. 1991. The nature of global 

change. Chapter 1. In: Walker BH, Steffen WL, Canadell 

J, Ingram J, editors. The Terrestrial Biosphere and Global 

Change. Cambridge, United Kingdom: Cambridge University 

Press. p 1-18. 

Type: Book Section

Geographic Area: Global

Compilers’ Keywords: climate change, land cover, atmospheric composition, biodiversity, synthesis, terrestrial 

ecosystems

Introduction: Despite the growing prominence of global 

change as an environmental issue, the term ‘global change’ 

and the phenomena it encompasses are still widely misunderstood. Global change is often referred to as ‘global warming’ or ‘climate change,’ but there is much more to it than a 

change in climate and, in fact, climatic change is now and 

will continue to be for the next few decades one of the less 

important components of global change in terms of effects 

on the structure and functioning of terrestrial ecosystems. 

Other more immediate and more certain components are 

the direct human conversion and modification of terrestrial 

CHAPTER 1 • CLIMATE CHANGE • GENERAL
12 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

36. Woodward FI. 1987. Climate and plant distribution. 

London, United Kingdom: Cambridge University Press. 188 p. 

Type: Book

Geographic Area: Global

Compilers’ Keywords: climate change, plant distribution, 

range shifts, migration

Introduction: The central thesis for plant ecology is that 

climate exerts the dominant control on the distribution of the 

major vegetation types of the world. Within a vegetation type 

smaller-scale variations in distribution may be controlled by 

smaller-scale features of the environment such as soil types, 

human activity or topography. However, at all spatial scales the 

response of the plant to climate is a crucial feature in its presence. In view of the importance of climate in controlling distribution of plants it is surprising that this area of subject is not 

a popular one in plant ecology. Such questions as ‘when’ and 

‘how’ does climate have its effect are clearly difficult to answer, 

perhaps explaining the limited acceptance as an area of study. I 

have set out to encourage an interest in climate and plant distribution by proposing different approaches to the subject. They 

are approaches which many not find general acceptance, but 

if they create interest and debate, which is my hope, then they 

will have been successful. I have drawn exclusively on vascular plants as examples of distribution types and responses. This 

reflects my interests and is not a comment on the ecology of 

non-vascular plants. Where possible, I have also avoided the 

use of the emotive terminologies, such as niche, strategy and 

adaptation; however I admit to the unavoidable use of competition, in spite of real problems in its measurements. 

Link: http://books.google.com/books?hl=en&lr=&id=0Ld1h0

MT3oIC&oi=fnd&pg=PR9&ots=QeJGze6gZK&sig=GtOVVn

kmqLYEBXlzL_cjCWLDBBY#v=onepage&q&f=false

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 13

37. Allen CD, Macalady AK, Chenchouni H, 

Bachelet D, McDowell N, Vennetier M, Kitzberger 

T, Rigling A, Breshears DD, Hogg EH, Gonzalez P, 

Fensham R, Zhang Z, Castro J, Demidova N, Lim 

J-H, Allard G, Running SW, Semerci A, Cobb NS. 

2010. A global overview of drought and heat-induced tree mortality reveals emerging climate change risks for forests. Forest 

Ecology and Management 259:660-684.

Type: Journal 

Geographic Area: Global

Keywords: climate change, drought effects, forest die-off, forest mortality, global patterns, tree mortality

Abstract: Greenhouse gas emissions have significantly altered global climate, and will continue to do so in the future. 

Increases in the frequency, duration, and/or severity of drought 

and heat stress associated with climate change could fundamentally alter the composition, structure, and biogeography 

of forests in many regions. Of particular concern are potential 

increases in tree mortality associated with climate induced 

physiological stress and interactions with other climate-mediated processes such as insect outbreaks and wildfire. Despite 

this risk, existing projections of tree mortality are based on 

models that lack functionally realistic mortality mechanisms, 

and there has been no attempt to track observations of climatedriven tree mortality globally. Here we present the first global 

assessment of recent tree mortality attributed to drought and 

heat stress. Although episodic mortality occurs in the absence 

of climate change, studies compiled here suggest that at least 

some of the world’s forested ecosystems already may be responding to climate change and raise concern that forests may 

become increasingly vulnerable to higher background tree 

mortality rates and die-off in response to future warming and 

drought, even in environments that are not normally considered water-limited. This further suggests risks to ecosystem 

services, including the loss of sequestered forest carbon and 

associated atmospheric feedbacks. Our review also identifies 

key information gaps and scientific uncertainties that currently hinder our ability to predict tree mortality in response 

to climate change and emphasizes the need for a globally coordinated observation system. Overall, our review reveals the 

potential for amplified tree mortality due to drought and heat 

in forests worldwide.

Link: http://static.consbio.org/media/publications/files/

Allen2010ForEcManGlobalForestDieback.pdf

General (1–36)—editorials, opinions, reviews, trends, surveys, summaries

Research (37–135)—climate change impacts, 

projections, paleoecology

Strategies (136–205)—policies, adaptation 

options, frameworks, mitigations

Resources (206–214)—tools, websites, software

Climate Change 

CHAPTER 1 • CLIMATE CHANGE • RESEARCH

#. Author(s), Date. Title. Source.

Type: 

Geographic Area: 

Keywords: 

Description:

Link:

location of study

keywords written by the 

author(s) or provided by the 

compilers

ID number and

information

abstract, summary, or 

portions of the introduction 

or conclusion written by the 

author(s) or provided by the 

compilers

book, book section, conference 

paper, dissertation, government 

document, journal article, 

newsletter, presentation, report, 

thesis or web page

internet address of reference

Each reference contains the following:
14 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

40. Barnosky AD, Hadly EA, Bascompte J, Berlow 

EL, Brown JH, Fortelius M, Getz WM, Harte J, 

Hastings A, Marquet PA, Martinez ND, Mooers A, 

Roopnarine P, Vermeij G, Williams JW, Gillespie R, 

Kitzes J, Marshall C, Matzke N, Mindell DP, Revilla 

E, Smith AB. 2012. Approaching a state shift in Earth’s biosphere. Nature 486:52-58.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: animals, climate change, ecosystem, 

environmental monitoring, forecasting, human activities, models

Abstract: Localized ecological systems are known to shift 

abruptly and irreversibly from one state to another when they 

are forced across critical thresholds. Here we review evidence 

that the global ecosystem as a whole can react in the same way 

and is approaching a planetary-scale critical transition as a result of human influence. The plausibility of a planetary-scale 

“tipping point” highlights the need to improve biological forecasting by detecting early warning signs of critical transitions 

on global as well as local scales, and by detecting feedbacks 

that promote such transitions. It is also necessary to address 

root causes of how humans are forcing biological changes.

Link: http://www.ncbi.nlm.nih.gov/pubmed/22678279

41. Beaulieu J, Rainville A. 2005. Adaptation to climate 

change: genetic variation is both a short- and a long-term solution.

The Forestry Chronicle 81:704-709.

Type: Journal 

Geographic Area: Eastern Canada

Keywords: climate change, white spruce, provenance test, 

transfer model, site index, adaptation, plantation, GIS 

Abstract: We propose a methodology combining a biophysical 

site index model and a seed source transfer model based on both 

temperature and precipitation to estimate white spruce plantation yield under present and future global warming conditions. 

The biophysical site index model predicts dominant height at 25 

years, which is further used to estimate plantation yield using 

yield tables. The transfer model shows that, on average, seed 

sources are best adapted to the temperature conditions where 

they presently grow, and give maximum yield under these conditions. However, this model also shows that transfer of seed 

sources to drier sites could improve plantation yield. To predict 

site index values under climate change conditions, values obtained from the biophysical site index model are corrected by a 

factor estimated using the seed source transfer model. Our simulation results predict that global warming should favor a slight 

increase in white spruce plantation yield in southern Quebec. 

However, one cannot expect to obtain similar yields from a seed 

source rapidly exposed to warmer conditions compared with a 

seed source that is presently growing under climatic conditions to 

which it has become adapted. It would take several generations 

38. Aubry C, Devine W, Shoal R, Bower AD, Miller 

J, Maggiulli N. 2011. Climate change and forest biodiversity: 

a vulnerability assessment and action plan for National Forests 

in Western Washington. Olympia (WA): USDA Forest Service. 

310 p.

Type: Government Document

Geographic Area: Pacific Northwest USA

Compilers’ Keywords: forest management, vulnerability 

assessment

Introduction: Climate change predictions for the Pacific 

Northwest include overall warming, increased winter precipitation, and decreased summer precipitation, resulting in warmer, 

wetter winters and warmer, drier summers. The extent and duration of the regional snowpack is projected to decrease, particularly at lower elevations. Seasonal stream flow patterns are 

likely to shift to earlier spring peak flows and lower summer 

flows, especially for snowmelt-dominated watersheds. There is 

a limited amount of information on climatic tolerance for many 

tree species and even less information on what complex interactions could result from ecosystem-wide exposure to a changing 

environment. The goals of this analysis are to conduct a climate 

change vulnerability assessment of forest tree species, assess 

the vulnerability of non-forested habitats to climate change, and 

propose practical management actions that will work under a 

variety of future climate scenarios and can be implemented by 

the national forests in western Washington in cooperation with 

other land managers. 

Link: http://ecoshare.info/2011/05/09/climate-change-andforest-biodiversity-a-vulnerability-assessment-and-action-planfor-national-forests-in-western-washington/

39. Bakkenes M, Eickhout B, Alkemade R. 2006.

Impacts of different climate stabilisation scenarios on plant species in Europe. Global Environmental Change 16:19-28.

Type: Journal 

Geographic Area: Europe

Keywords: biodiversity, climate change, stabilisation profiles, 

integrated assessment, scenario analysis

Abstract: With the use of goals from the Convention on 

Biological Diversity we evaluated two climate stabilisation profiles on their merits for conservation of biodiversity, comparing 

them with a baseline profile. Focusing on plant ecosystems at 

the pan-European level, we concluded that although a maximum 

global-mean temperature increase of 21°C is likely to be met 

in a 550 ppmv CO2-equivalent stabilisation profile, large areas 

of ecosystems in Europe will be affected. Most of the impacts 

manifest themselves in northern countries, with a high net increase of plant species, and in Mediterranean countries, with a 

decrease in the number of plant species and stable area. Other 

impacts are less robust, given the regional variation in climate 

results for different climate models.

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 15

origin. Therefore, we recommend enhancing the genetic variability within populations of species in general.

Link: http://www.researchgate.net/publication/224763331_

Ecotypes_of_European_grass_species_respond_specifically_

to_warming_and_extreme_drought/file/72e7e51a5c38a6cf72.

pdf

43. Bell G, Gonzalez A. 2011. Adaptation and evolutionary rescue in metapopulations experiencing environmental deterioration. Science 332:1327-1330.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: climate change, physiological, biological evolution, models, directed molecular evolution, adaptation, 

Saccharomyces cerevisiae

Abstract: It is not known whether evolution will usually 

be rapid enough to allow a species to adapt and persist in a 

deteriorating environment. We tracked the eco-evolutionary dynamics of metapopulations with a laboratory model 

system of yeast exposed to salt stress. Metapopulations 

experienced environmental deterioration at three different 

rates and their component populations were either unconnected or connected by local dispersal or by global dispersal. 

We found that adaptation was favored by gradual deterioration 

and local dispersal. After further abrupt deterioration, the frequency of evolutionary rescue depended on both the prior rate 

of deterioration and the rate of dispersal. Adaptation was surprisingly frequent and rapid in small peripheral populations. Thus, 

evolutionary dynamics affect both the persistence and the 

range of a species after environmental deterioration.

Link: http://izt.ciens.ucv.ve/ecologia/Archivos/ECO_POB%20

2011/ECOPO2_2011/Bell%20y%20Gonzalez%202011.pdf

44. Bentz B, editor. 2009. Bark beetle outbreaks in western North America: causes and consequences. Snowbird (UT): 

University of Utah Press. 42 p. 

Type: Conference Proceedings

Geographic Area: Western North America

Compilers’ Keywords: climate change, forest impact, insect 

outbreaks, symposium

Compilers’ Summary: Since 1990, native bark beetles have 

killed billions of trees across millions of acres of forest from 

Alaska to northern Mexico. Although bark beetle infestations 

are a regular force of natural change in forested ecosystems, 

several of the current outbreaks, which are occurring simultaneously across western North America, are the largest and most 

severe in recorded history.

Link: http://www.usu.edu/beetle/Symposium/

SymposiumCover.pdf

(adaptation lag) for a seed source to adapt to warmer conditions 

to which it has become adapted. We believe that the method we 

propose will be helpful in identifying the most productive seed 

source to be used at any given location in the province, and in 

revising seed source transfer rules. 

Link: http://mrn.gouv.qc.ca/publications/forets/connaissances/

recherche/Rainville-Andre/For-Chron-81-5-704-709.pdf

42. Beierkuhnlein C, Thiel D, Jentsch A, Willner E, 

Kreyling J. 2011. Ecotypes of European grass species respond 

differently to warming and extreme drought. Journal of Ecology 

99:703-713.

Type: Journal 

Geographic Area: Europe

Keywords: adaptation, climate change, climatic events, extreme weather, grassland, meadow, phenotypes, plant-climate 

interactions, provenances, within-species vulnerability

Abstract: Climate extremes are expected to increase in frequency and magnitude as a consequence of global warming. 

Managed permanent grasslands cover a large surface in Europe 

and contribute substantially to agricultural production. These 

managed plant communities are dominated by perennial clonal 

species. Their capacity to adapt to rapidly changing environmental conditions may be limited. We hypothesize that those plant 

populations that have already been exposed to conditions that 

are expected to occur due to future climate change, particularly 

conditions that would be “extreme” in the target area, are able 

to cope better with these conditions. For a common-garden 

experiment we selected ecotypes (provenances as supported 

by accessions in seed banks) of important European grass 

species: Arrhenatherum elatius, Festuca pratensis, Holcus lanatus and Alopecurus pratensis. Southern target locations of 

ecotypes (populations) were identified based on climate model 

projections for the local site in Northern Bavaria, Germany. In a 

controlled experiment, the plants were exposed to warming and 

extreme drought. Drought conditions (16–19 days, depending 

on the species) were imposed starting from the end of May in 

combination with and without an increase in the average temperature from May to September 2009 (+1.5 °K compared with 

control; +2.5 °K compared with ambient conditions outside of 

the experimental units). Ecotypes and drought manipulation had 

significant impacts on biomass production and tissue die-back. 

Significant interactions between ecotype and drought indicated 

a different drought tolerance of the ecotypes in some cases. The 

warming treatment yielded a less significant response. The local 

ecotype generally did not perform significantly worse than the 

presumably better-adapted southern ecotypes. The selection of 

ecotypes that are adapted to more extreme climatic conditions 

could be an option for maintaining future ecosystem functioning 

in temperate managed grasslands, as was indicated by the clear 

differences between ecotypes in our experiment. Based on our 

data, however, performance cannot be predicted from climatic 

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
16 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

the vulnerability of both farmers and species is dependent on the scenario under consideration. In agriculture, 

it is the socio-economic scenarios that particularly lead 

to different patterns of intensification, extensification and 

abandonment. For species, vulnerability is more related to 

the climate change scenarios. In both cases, the adaptation options and potential were associated with the different socio-economic futures and policy intervention. The 

conceptual Linking of the two sectors shows that impacts 

in the agriculture sector and consequent adaptation could 

have a significant effect on the adaptation potential of species. This demonstrates the importance of cross-sectoral 

assessments of vulnerability and highlights the importance of sectoral integration in policy development and 

implementation.

Link: http://ethree.com/downloads/Climate%20Change%20

Readings/International%20Climate%20Policy/Berry%20-%20

Vulnerability%20Agr%20land%20%2B%20species%20climate%20change.pdf

47. Booth TH, Broadhurst LM, Pinkard E, Prober 

SM, Dillon SK, Bush D, Pinyopusarerk K, Doran JC, 

Ivkovich M, Young AG. 2015. Native forests and climate 

change: lessons from eucalypts. Forest Ecology and Management 

347:18-29.

Type: Journal 

Geographic Area: Australia

Keywords: vulnerability, adaptation, carbon dioxide, genomics, species distribution models, simulation models

Abstract: The purpose of this paper is to review studies relevant to potential climate change impacts on natural stands of 

eucalypts, with a view to identifying not only specific lessons 

for the management of native forests in Australia but also some 

general lessons relevant to native forests anywhere. More than 

800 species of Eucalyptus are found naturally across Australia, 

as well as species such as E. deglupta and E. urophylla in countries north of Australia. Eucalypts provide a particularly interesting opportunity to examine the likely impacts of climate change, 

as many species have been widely evaluated in trials within and 

outside Australia, often under conditions that are warmer and 

sometimes drier than those found within their natural distributions. Results from these trials indicate the intrinsic ability of 

particular eucalypt species and provenances to tolerate conditions that are somewhat different from those experienced within 

their natural distributions. Eucalypts have particularly poor 

dispersal capabilities, so natural stands will be generally unable 

to track changing climatic conditions. Therefore, in the period 

to the end of the present century a key issue for each eucalypt 

species under climate change is whether its intrinsic adaptability will be sufficient to allow it to survive where it is currently 

located. Their ability to survive will be affected not only by climatic, but also atmospheric changes, which will affect important 

45. Bentz BJ, Régnière J, Fettig CJ, Hansen 

EM, Hayes JL, Hicke JA, Kelsey RG, Negrón JF, 

Seybold SJ. 2010. Climate change and bark beetles of the 

western United States and Canada: direct and indirect effects.

BioScience 60:602-613.

Type: Journal 

Geographic Area: Western North America

Keywords: cold tolerance, mountain pine beetle, seasonality, 

spruce beetle, temperature 

Abstract: Climatic changes are predicted to significantly affect the frequency and severity of disturbances that shape forest 

ecosystems. We provide a synthesis of climate change effects 

on native bark beetles, important mortality agents of conifers in 

western North America. Because of differences in temperaturedependent life-history strategies, including cold-induced mortality and developmental timing, responses to warming will differ 

among and within bark beetle species. The success of bark beetle populations will also be influenced indirectly by the effects of 

climate on community associates and host-tree vigor, although 

little information is available to quantify these relationships. We 

used available population models and climate forecasts to explore the responses of two eruptive bark beetle species. Based 

on projected warming, increases in thermal regimes conducive 

to population success are predicted for Dendroctonus rufipennis

(Kirby) and Dendroctonus ponderosae Hopkins, although there is 

considerable spatial and temporal variability. These predictions 

from population models suggest a movement of temperature 

suitability to higher latitudes and elevations and identify regions 

with a high potential for bark beetle outbreaks and associated 

tree mortality in the coming century.

Link: http://www.fs.fed.us/wwetac/projects/PDFs/BioScience_

CC_and_Bark_Beetles.pdf

46. Berry PM, Rounsevell MDA, Harrison PA, 

Audsley E. 2006. Assessing the vulnerability of agricultural 

land use and species to climate change and the role of policy 

in facilitating adaptation. Environmental Science & Policy 

9:189-204.

Type: Journal 

Geographic Area: Europe

Keywords: climate change, farmers, species, vulnerability, adaptation, policy

Abstract: The term vulnerability has been used in a variety of contexts, including climate change impact assessment. In this paper those issues relevant to climate change 

impacts on agriculture and species are discussed. Outputs 

from models are used to assess the vulnerability of farmers and species to climate and socio-economic change by 

estimating their sensitivity and capacity to adapt to external factors as a means of identifying what causes the 

differences in their vulnerability. The results showed that 

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 17

49. Breshears DD, Cobb NS, Rich PM, Price KP, Allen 

CD, Balice RG, Romme WH, Kastens JH, Floyd ML, 

Belnap J, Anderson JJ, Myers OB, Meyer CW. 

2005. Regional vegetation die-off in response to global-changetype drought. Proceedings of the National Academy of Science 

USA 102:15144-15148.

Type: Journal 

Geographic Area: Southwestern USA 

Keywords: tree mortality, vegetation dynamics, climate change 

impacts, woodlands, Pinus edulis

Abstract: Future drought is projected to occur under warmer 

temperature conditions as climate change progresses, referred 

to here as global-change-type drought, yet quantitative assessments of the triggers and potential extent of drought-induced 

vegetation die-off remain pivotal uncertainties in assessing 

climate-change impacts. Of particular concern is regional-scale 

mortality of overstory trees, which rapidly alters ecosystem type, 

associated ecosystem properties, and land surface conditions for 

decades. Here, we quantify regional-scale vegetation die-off 

across southwestern North American woodlands in 2002-2003 

in response to drought and associated bark beetle infestations. 

At an intensively studied site within the region, we quantified 

that after 15 months of depleted soil water content, > 90% of 

the dominant, overstory tree species (Pinus edulis, a pinon) died. 

The die-off was reflected in changes in a remotely sensed index of vegetation greenness (Normalized Difference Vegetation 

Index), not only at the intensively studied site but also across 

the region, extending over 12,000 km2 or more; aerial and field 

surveys confirmed the general extent of the die-off. Notably, the 

recent drought was warmer than the previous subcontinental 

drought of the 1950s. The limited, available observations suggest 

that die-off from the recent drought was more extensive than that 

from the previous drought, extending into wetter sites within the 

tree species’ distribution. Our results quantify a trigger leading 

to rapid, drought-induced die-off of overstory woody plants at 

subcontinental scale and highlight the potential for such die-off 

to be more severe and extensive for future global-change-type 

drought under warmer conditions.

Link: http://www.ncbi.nlm.nih.gov/pubmed/16217022

50. Bucharová A, Brabec J, Münzbergová Z. 

2012. Effect of land use and climate change on the future 

fate of populations of an endemic species in central Europe.

Biological Conservation 145:39-47.

Type: Journal 

Geographic Area: Central Europe

Keywords: conservation action plan, population viability analysis, grassland management, Gentianella, matrix model, extinction probability

Abstract: The identification of optimal management strategies 

for a given species is a major challenge of species conservation. 

processes such as photosynthesis and water exchange. Again eucalypts provide a useful group for climate change studies as their 

commercial significance has led to various enhanced carbon dioxide experiments being carried out, as well as detailed genomic 

studies. This review considers eucalypts in relation to four main 

areas: (i) resources and characteristics (natural distributions and 

introduced distributions including their adaptability/plasticity), 

(ii) analysis tools (species distribution models and growth models), (iii) physiological factors (including temperature, drought 

and enhanced CO2) and (iv) interactions with other species 

(including pests and diseases). Priorities for future research are 

identified. It is concluded that analyses that do not allow for the 

intrinsic climatic adaptability of tree species, as well as their 

particular dispersal capabilities, are unlikely to provide reliable 

predictions of climate change impacts.

Link: http://www.sciencedirect.com/science/article/pii/

S0378112715001139

48. Bradley BA, Wilcove DS. 2009. When invasive plants 

disappear: transformative restoration possibilities in the western 

United States resulting from climate change. Restoration Ecology 

17:715-721.

Type: Journal 

Geographic Area: Western USA

Keywords: bioclimatic envelope modeling, Bromus tectorum, 

climate change, ecological niche, invasive species, restoration, 

species distribution

Abstract: Most ecologists believe that climate change 

poses a significant threat to the persistence of native species. 

However, in some areas climate change may reduce or eliminate nonnative invasive species, creating opportunities for 

restoration. If invasive species are no longer suited to novel 

climate conditions, the native communities that they replaced 

may not be viable either. If neither invasive nor native species 

are climatically viable, a type of “transformative” restoration 

will be required, involving the translocation of novel species 

that can survive and reproduce under new climate conditions. 

Here, we illustrate one approach for restoration planning by 

using bioclimatic envelope modeling to identify restoration 

opportunities in the western United States, where the invasive 

plant cheatgrass (Bromus tectorum) is no longer climatically 

viable under 2100 conditions projected by the Geophysical 

Fluid Dynamics Laboratory (GFDL2.1) coupled atmosphereocean general circulation model. We then select one example 

of a restoration target area and identify novel plant species 

that could become viable at the site in the wake of climate 

change. We do so by identifying the closest sites that currently 

have climate conditions similar to those projected at the restoration target area in 2100. This approach is a first step toward 

identifying appropriate species for transformative restoration.

Link: http://www.firescience.gov/projects/05-2-1-94/project/05-2-1-94_bradley_re_2009.pdf

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
18 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

the Iberian Peninsula region. We found that defoliation trends 

are paralleled by significant increases in tree mortality rates in 

drier areas that are related to tree density and temperature effects. Furthermore, we show that severe drought impacts are 

associated with sudden changes in insect and fungal defoliation 

dynamics, creating long-term disruptive effects of drought on 

food webs. Our results reveal a complex geographical mosaic 

of species-specific responses to climate change—driven drought 

pressures on the Iberian Peninsula, with an overwhelmingly predominant trend toward increased drought damage.

Link: http://www.pnas.org/content/108/4/1474.short

52. Chmura DJ, Anderson PD, Howe GT, Harrington 

CA, Halofsky JE, Peterson DL, Shaw DC, St. Clair JB. 

2011. Forest responses to climate change in the northwestern 

United States: ecophysiological foundations for adaptive management. Forest Ecology and Management 261:1121-1142.

Type: Journal 

Geographic Area: Northwestern USA 

Keywords: adaptation, drought, fire, genetics, insects, 

silviculture

Abstract: Climate change resulting from increased concentrations of atmospheric carbon dioxide ([CO2]) is expected to result in warmer temperatures and changed precipitation regimes 

during this century. In the northwestern US, these changes will 

likely decrease snowpack, cause earlier snowmelt, increase 

summer evapotranspiration, and increase the frequency and 

severity of droughts. Elevated [CO2] and warmer temperatures 

may have positive effects on growth and productivity where 

there is adequate moisture or growth is currently limited by cold. 

However, the effects of climate change are generally expected 

to reduce growth and survival, predispose forests to disturbance 

by wildfire, insects, and disease; and ultimately change forest 

structure and composition at the landscape scale. Substantial 

warming will likely decrease winter chilling resulting in delayed 

bud burst, and adversely affect flowering and seed germination 

for some species. The extent of these effects will depend on the 

magnitude of climate change, the abilities of individual trees to 

acclimate, and for tree populations to adapt in situ, or to migrate 

to suitable habitats. These coping mechanisms may be insufficient to maintain optimal fitness of tree populations to rapidly 

changing climate. Physiological responses to climatic stresses 

are relatively well-understood at the organ or whole-plant scale 

but not at the stand or landscape scale. In particular, the interactive effects of multiple stressors are not well known. Genetic and 

silvicultural approaches to increase adaptive capacities and to 

decrease climate-related vulnerabilities of forests can be based 

on ecophysiological knowledge. Effective approaches to climate 

adaptation will likely include assisted migration of species and 

populations, and density management. Use of these approaches to increase forest resistance and resilience at the landscape 

scale requires a better understanding of species adaptations, 

It becomes especially challenging when the environmental 

conditions are expected to change in the future, and the optimal management applied today may differ from the management that is optimal under the changed conditions (e.g. due to 

climate change). This study evaluates prospect of a rare plant 

species endemic to semi-natural grasslands in central Europe, 

Gentianella praecox subsp. bohemica. The number of populations of this species has declined rapidly in the last 60 years; 

currently, a conservation action plan has been established in 

the Czech Republic, where most populations of this species 

occur. This study uses periodic matrix models to compare different management regimes under different scenarios of climate 

change and to identify the optimal management in each case. 

Without management, populations of the species are not able to 

survive. Flowering individuals can occur for a long time after 

the cessation of management, but the extinction of the population is inevitable within several decades. Without management, 

even very large populations (1,000 flowering individuals) will 

go extinct in less than 50 years. Total extinction (including seed 

bank) will follow several years after observation of the last 

flowering plant. The most suitable management is mowing and 

disturbance (by harrowing), which is also the best method for 

restoration of threatened populations. Mowing is less suitable, 

but it is fully sufficient for large prospering populations. When 

managed, even small populations (10–15 flowering individuals) 

are able to survive. When management is applied, future climate 

change may have a relatively small impact on the probability of 

survival of the species. Climate change will, however, increase 

the extinction probability of very small populations.

51. Carnicer J, Coll M, Ninyerola M, Pons X, 

Sanchez G, Penuelas J. 2011. Widespread crown condition decline, food web disruption, and amplified tree mortality 

with increased climate change-type drought. Proceedings of the 

National Academy of Science USA 108:1474-1478.

Type: Journal 

Geographic Area: Southern Europe

Keywords: extreme events, earth system feedbacks, ecological 

networks, global change, Mediterranean biome

Abstract: Climate change is progressively increasing severe 

drought events in the Northern Hemisphere, causing regional 

tree die-off events and contributing to the global reduction of 

the carbon sink efficiency of forests. There is a critical lack of 

integrated community-wide assessments of drought-induced 

responses in forests at the macroecological scale, including defoliation, mortality, and food web responses. Here we report a 

generalized increase in crown defoliation in southern European 

forests occurring during 1987–2007. Forest tree species have 

consistently and significantly altered their crown leaf structures, 

with increased percentages of defoliation in the drier parts of 

their distributions in response to increased water deficit. We assessed the demographic responses of trees associated with increased defoliation in southern European forests, specifically in 

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 19

how a half-dozen tree species differ in their responses to monthly variation in four climatic-related variables: water availability, 

deviations from an optimum temperature, atmospheric humidity deficits, and the frequency of frost. Rather than use climatic 

data directly to correlate with a species’ distribution, we assess 

the relative constraints of each of the four variables as they affect predicted monthly photosynthesis for Douglas-fir, the most 

widely distributed species in the region. We apply an automated 

regression-tree analysis to create a suite of rules, which differentially rank the relative importance of the four climatic modifiers 

for each species, and provide a basis for predicting a species’ 

presence or absence on 3,737 uniformly distributed U.S. Forest 

Services’ Forest Inventory and Analysis (FIA) field survey plots. 

Results of this generalized rule-based approach were encouraging, with weighted accuracy, which combines the correct prediction of both presence and absence on FIA survey plots, averaging 87%. A wider sampling of climatic conditions throughout 

the full range of a species’ distribution should improve the basis 

for creating rules and the possibility of predicting future shifts in 

the geographic distribution of species.

Link: http://www.fs.fed.us/rm/pubs_other/rmrs_2009_coops_

n001.pdf

55. Cortini F, Comeau PG, Bokalo M. 2012. Trembling 

aspen competition and climate effects on white spruce growth 

in boreal mixtures of Western Canada. Forest Ecology and 

Management 277:67-73.

Type: Journal 

Geographic Area: Western Canada

Keywords: conifer growth, mixedwood forests, climate, competition, white spruce, trembling aspen

Abstract: We investigated the combined effect of trembling aspen competition and climate on white spruce growth using data 

from a long term study with matching treatments spread across 

the boreal mixed wood forests of Alberta and Saskatchewan 

(Canada). Results indicate that competition (i.e., aspen basal 

area), initial size of the tree and mean annual temperature can 

account for 88% of the year to year variation in spruce volume 

growth for these six locations. Based on the model that we developed, spruce growth, in the absence of competition, is estimated 

to increase by up to 17% compared with an increase in mean 

annual temperature from 2 °C to 3.3 °C, while at high levels 

of competition (aspen basal area = 27 m2/ha) spruce growth increases by only 8%. Moreover, effects of aspen on spruce growth 

increase more than proportionally as temperature increases. 

This outcome indicates that abundant aspen competition limits 

the spruce responses to rising temperature, presumably due to 

competition for light and potentially increased competition for 

soil resources. Results also show that competition and climate 

effects vary between locations, indicating that spruce growth is 

strongly influenced by local factors such as micro-climate, topography, and soil properties. 

within-species genetic variation, and the mitigating effects of 

silvicultural treatments.

Link: http://www.treesearch.fs.fed.us/pubs/39611

53. Classen AT, Norby RJ, Campany CE, Sides KE, 

Weltzin JF. 2010. Climate change alters seedling emergence and establishment in an old-field ecosystem. PLoS ONE 

5:e13476.

Type: Journal 

Geographic Area: Eastern USA

Compilers’ Keywords: atmosphere, carbon dioxide, tree 

growth and development

Abstract: Ecological succession drives large-scale changes in 

ecosystem composition over time, but the mechanisms whereby climatic change might alter succession remain unresolved. 

Here, we asked if the effects of atmospheric and climatic change 

would alter tree seedling emergence and establishment in an oldfield ecosystem, recognizing that small shifts in rates of seedling emergence and establishment of different species may have 

long-term repercussions on the transition of fields to forests in 

the future. We introduced seeds from three early successional 

tree species into constructed old-field plant communities that had 

been subjected for 4 years to altered temperature, precipitation, 

and atmospheric CO2 regimes in an experimental facility. Our 

experiment revealed that different combinations of atmospheric 

CO2 concentration, air temperature, and soil moisture altered 

seedling emergence and establishment. Treatments directly and 

indirectly affected soil moisture, which was the best predictor of 

seedling establishment, though treatment effects differed among 

species. The observed impacts, coupled with variations in the 

timing of seed arrival, are demonstrated as predictors of seedling emergence and establishment in ecosystems under global 

change.

Link: http://www.ncbi.nlm.nih.gov/pubmed/20976104

54. Coops NC, Waring RH, Schroeder TA. 2009. 

Combining a generic process-based productivity model and a 

statistical classification method to predict the presence and absence of tree species in the Pacific Northwest, USA. Ecological 

Modelling 220:1787-1796.

Type: Journal 

Geographic Area: Pacific Northwest USA

Keywords: 3-PG model, regression-tree analysis, climate 

change, U.S. Forest Inventory and Analysis, Sitka spruce, ponderosa pine, western juniper, lodgepole pine, Douglas-fir, western hemlock

Abstract: Although long-lived tree species experience considerable environmental variation over their life spans, their 

geographical distributions reflect sensitivity mainly to mean 

monthly climatic conditions. We introduce an approach that 

incorporates a physiologically based growth model to illustrate 

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
20 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

geographic area of interest, and possibly on what climate projections are available for a region. Rather than a misguided attempt to identify the “most accurate” climate scenario, managers are strongly encouraged to explore variability through the 

use of multiple climate scenarios. Considering a range of possible future climates facilitates the identification of management 

strategies to help ensure resilience of natural resource systems 

across a broad set of potential conditions. Downscaling climate 

projections increases the spatial resolution of climate information and can make projections more relevant to natural resource 

managers by allowing decision-makers to better visualize what 

these different futures imply locally and regionally. The following series of questions describes key concepts that end-users of 

climate projection products should understand to appropriately 

interpret downscaled climate projections, including various 

sources of uncertainty. The selection used for each component of 

a downscaled climate projection has implications for interpreting the resulting climate scenario. Understanding the merits and 

limitations of the downscaling method employed is also important since downscaling approaches vary in their dependence on 

observed data availability, computational requirements, and in 

resultant uncertainty owed to biases of the method or the spatial 

scale of the downscaling.

Link: http://www.treesearch.fs.fed.us/pubs/40614

58. Davison JE, Coe S, Finch DM, Rowland EL, 

Friggens MM, Graumlich LJ. 2012. Bringing indices of 

species vulnerability to climate change into geographic space: an 

assessment across the Coronado National Forest. Biodiversity 

and Conservation 21:189-204.

Type: Journal 

Geographic Area: Southwestern USA

Keywords: biodiversity, climate change adaptation, GIS, land 

use planning, species vulnerability assessments

Abstract: Indices that rate the vulnerability of species to climate change in a given area are increasingly used to inform 

conservation and climate change adaptation strategies. These 

species vulnerability indices (SVI) are not commonly associated 

with landscape features that may affect local-scale vulnerability. 

To do so would increase their utility by allowing managers to 

examine how the distributions of vulnerable species coincide 

with environmental features such as topography and land use, 

and to detect landscape-scale patterns of vulnerability across 

species. In this study we evaluated 15 animal species that had 

been scored with the USDA-Forest Service Rocky Mountain 

Research Station’s system for assessing vulnerability of species 

to climate change. We applied the vulnerability scores to each 

species’ respective habitat models in order to visualize the spatial patterns of cross-species vulnerability across the biologically 

diverse Coronado National Forest, and to identify the considerations of spatially referencing such indices. Across the study extent, cross-species vulnerability was higher in higher-elevation 

Link: http://www.researchgate.net/publication/225071346_

Trembling_aspen_competition_and_climate_effects_on_

white_spruce_growth_in_boreal_mixtures_of_Western_

Canada/file/9fcfd4fc66733d4533.pdf

56. Crookston NL, Rehfeldt GE, Dixon GE, 

Weiskittel AR. 2010. Addressing climate change in the forest vegetation simulator to assess impacts on landscape forest 

dynamics. Forest Ecology and Management 260:1198-1211.

Type: Journal 

Geographic Area: Western USA

Keywords: species–climate relationships, stand dynamic, species composition, genetic adaptation, general circulation model, 

climate change, carbon loads, site index, growth and yield

Abstract: To simulate stand-level impacts of climate change, 

predictors in the widely used Forest Vegetation Simulator (FVS) 

were adjusted to account for expected climate effects. This 

was accomplished by: (1) adding functions that Link mortality 

and regeneration of species to climate variables expressing climatic suitability, (2) constructing a function Linking site index 

to climate and using it to modify growth rates, and (3) adding 

functions accounting for changing growth rates due to climateinduced genetic responses. For three climatically diverse landscapes, simulations were used to explore the change in species 

composition and tree growth that should accompany climate 

change during the 21st century. The simulations illustrated the 

changes in forest composition that could accompany climate 

change. Projections were the most sensitive to mortality, as the 

loss of trees of a dominant species heavily influenced stand dynamics. While additional work is needed on fundamental plant–

climate relationships, this work incorporates climatic effects into 

FVS to produce a new model called Climate–FVS. This model 

provides for managers a tool that allows climate change impacts 

to be incorporated in forest plans.

Link: http://www.treesearch.fs.fed.us/pubs/37335

57. Daniels AE, Morrison JF, Joyce LA, Crookston NL, 

Chen SC, McNulty SG. 2012. Climate projections FAQ.

Fort Collins (CO): USDA Forest Service, Rocky Mountain 

Research Station. General Technical Report RMRS-GTR277WWW. 32 p.

Type: Government Document

Geographic Area: Western USA

Keywords: climate change, climate projections, downscaling, 

general circulation models, vulnerability assessment

Abstract: Climate scenarios offer one way to identify and 

examine the land management challenges posed by climate 

change. Selecting projections, however, requires careful consideration of the natural resources under study, and where and 

how they are sensitive to climate. Selection also depends on 

the robustness of different projections for the resources and 

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 21

structures and functions (resilience), is likely to vary across environmental gradients such as moisture availability, but has received little study. We examined conifer regeneration a decade 

following complete stand-replacing wildfire in dry coniferous 

forests spanning a 700 m elevation gradient where low elevation 

sites had relatively high moisture stress due to the combination 

of high temperature and low precipitation. Conifer regeneration varied strongly across the elevation gradient, with little tree 

regeneration at warm and dry low elevation sites. Logistic regression models predicted rapid increases in regeneration across 

the elevation gradient for both seedlings of all conifer species 

and ponderosa pine seedlings individually. This pattern was especially pronounced for well-established seedlings (≥ 38 cm in 

height). Graminoids dominated lower elevation sites following 

wildfire, which may have added to moisture stress for seedlings 

due to competition for water. These results suggest moisture 

stress can be a critical factor limiting conifer regeneration following stand-replacing wildfire in dry coniferous forests, with 

predicted increases in temperature and drought in the coming 

century likely to increase the importance of moisture stress. 

Strongly moisture limited forested sites may fail to regenerate 

for extended periods after stand-replacing disturbance, suggesting these sites are high priorities for management intervention 

where maintaining forests is a priority.

61. Dukes JS, Pontius J, Orwig D, Garnas JR, Rodgers 

ViL, Brazee N, Cooke B, Theoharides KA, Stange 

EE, Harrington R, Ehrenfeld J, Gurevitch J, Lerdau 

M, Stinson K, Wick R, Ayres M. 2009. Responses of 

insect pests, pathogens, and invasive plant species to climate 

change in the forests of northeastern North America: what can 

we predict? Canadian Journal of Forest Research 39:231-248.

Type: Journal 

Geographic Area: Eastern North America

Compilers’ Keywords: uncertainty, management, USA, 

Canada

Abstract: Climate models project that by 2100, the northeastern U.S. and eastern Canada will warm by approximately 3-5 

°C with increased winter precipitation. These changes will affect 

trees directly and also indirectly through effects on “nuisance” 

species, such as insect pests, pathogens, and invasive plants. We 

review how basic ecological principles can be used to predict 

nuisance species’ responses to climate change and how this is 

likely to impact northeastern forests. We then examine in detail the potential response of two pest species (hemlock woolly 

adelgid (Adeleges tsugae Annand) and forest tent caterpillar 

(Malacosoma disstria Hubner)); two pathogens (armillaria root 

rot (Armillaria spp.) and beech bark disease (Cryptococcus fagisuga Lind. + Neonectria spp.)); and two invasive plant species (glossy buckthorn (Frangula alnus Mill.) and oriental 

bittersweet (Celastrus orbiculatus Thunb.)). Several of these 

species are likely to have stronger or more widespread effects 

on forest composition and structure under the projected climate. 

woodlands and lower in desert scrub. The results of spatially 

referencing SVI scores may vary according to the species examined, the area of interest, the selection of habitat models, and 

the method by which cross-species vulnerability indices are created. We show that it is simple and constructive to bring species 

vulnerability indices into geographic space: landscape-scale patterns of vulnerability can be detected, and relevant ecological 

and socioeconomic contexts can be taken into account, allowing 

for more robust conservation and management strategies.

Link: http://www.fs.fed.us/rm/pubs_other/rmrs_2012_davison_j001.pdf

59. Dawson TP, Jackson ST, House JI, Prentice IC, 

Mace GM. 2011. Beyond predictions: biodiversity conservation in a changing climate. Science 332:53-58.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: conservation of natural resources, migration, review, vulnerability, paleoecology, framework 

Abstract: Climate change is predicted to become a major threat 

to biodiversity in the 21st century, but accurate predictions and 

effective solutions have proved difficult to formulate. Alarming 

predictions have come from a rather narrow methodological 

base; but a new, integrated science of climate-change biodiversity assessment is emerging, based on multiple sources and 

approaches. Drawing on evidence from paleoecological observations, recent phenological and microevolutionary responses, 

experiments, and computational models, we review the insights 

that different approaches bring to anticipating and managing the 

biodiversity consequences of climate change, including the extent of species’ natural resilience. We introduce a framework that 

uses information from different sources to identify vulnerability 

and to support the design of conservation responses. Although 

much of the information reviewed is on species, our framework 

and conclusions are also applicable to ecosystems, habitats, ecological communities, and genetic diversity, whether terrestrial, 

marine, or fresh water.

Link: http://www.ncbi.nlm.nih.gov/pubmed/21454781

60. Dodson EK, Root HT. 2013. Conifer regeneration following stand-replacing wildfire varies along an elevation gradient in a ponderosa pine forest, Oregon, USA. Forest Ecology 

and Management 302:163-170.

Type: Journal 

Geographic Area: Oregon, USA

Keywords: resilience, environmental gradient, climate changetype drought, seedling establishment, natural regeneration

Abstract: Climate change is expected to increase disturbances 

such as stand-replacing wildfire in many ecosystems, which 

have the potential to drive rapid turnover in ecological communities. Ecosystem recovery, and therefore maintenance of critical 

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
22 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

potentially altering integrated suites of life-history traits. We 

examined evolutionary change in multiple life history traits 

of the annual plant Brassica rapa collected before and after 

a recent 5-year drought in southern California. We used 

a direct approach to examining evolutionary change by 

comparing ancestors and descendants. Collections were 

made from two populations varying in average soil moisture levels, and lines propagated from the collected seeds 

were grown in a greenhouse and experimentally subjected to 

conditions simulating either drought (short growing season) 

or high precipitation (long growing season) years. Comparing 

ancestors and descendants, we found that the drought caused 

many changes in life history traits, including a shift to earlier 

flowering, longer duration of flowering, reduced peak flowering and greater skew of the flowering schedule. Descendants 

had thinner stems and fewer leaf nodes at the time of flowering than ancestors, indicating that the drought selected for plants 

that flowered at a smaller size and earlier ontogenetic stage rather than selecting for plants to develop more rapidly. Thus, there 

was no evidence for absolute developmental constraints to flowering time evolution. Common principal component analyses 

showed substantial differences in the matrix of trait covariances 

both between short and long growing season treatments and 

between populations. Although the covariances matrices were 

generally similar between ancestors and descendants, there was 

evidence for complex evolutionary changes in the relationships 

among the traits, and these changes depended on the population 

and treatment. These results show that a full appreciation of the 

impacts of global change on phenotypic evolution will entail an 

understanding of how changes in climatic conditions affect trait 

values and the structure of relationships among traits.

64. Grace J. 1987. Climate tolerance and the distribution of 

plants. New Phytologist 106:113-130.

Type: Journal 

Geographic Area: Global

Keywords: plant distribution, tree line, life forms, temperature 

climatological variables

Summary: Boundaries to the distribution of life forms and species often coincide with isometric lines of climatological variables. Prominent examples of this phenomenon are the restriction of megaphanerophytes to areas where the mean temperature 

of the warmest month does not fall below about 10 °C, and the 

restriction of species within the British Isles to areas of specific 

summer or winter warmth. In the interpretation of these patterns, it is important to realize that the climate at plant surfaces 

may differ appreciably from that of the atmosphere as a whole, 

to an extent which depends on structural attributes of the vegetation. In particular, chamaephytes experience a much higher 

surface temperature than phanerophytes when comparisons are 

made in bright sunlight and this contributes to their success in 

mountains. Temperatures of flowers may similarly be elevated. 

In the British Isles, temperatures in the summer months may be 

However, uncertainty pervades our predictions because we lack 

adequate data on the species and because some species depend 

on complex, incompletely understood, unstable relationships. 

While targeted research will increase our confidence in making 

predictions, some uncertainty will always persist. Therefore, we 

encourage policies that allow for this uncertainty by considering 

a wide range of possible scenarios. 

Link: http://web.ics.purdue.edu/~jsdukes/DukesEtAl2009.pdf

62. Finch DM. 2012. Climate change in grasslands, shrublands, and deserts of the interior American West: a review and 

needs assessment. Fort Collins (CO): USDA Forest Service, 

Rocky Mountain Research Station. General Technical Report 

RMRS-GTR-285. 139 p.

Type: Government Document

Geographic Area: Western USA

Keywords: climate change, grasslands, shrublands, deserts, 

assessment

Abstract: Recent research and species distribution modeling predict large changes in the distributions of species and 

vegetation types in the western interior of the United States 

in response to climate change. This volume reviews existing 

climate models that predict species and vegetation changes 

in the western United States, and it synthesizes knowledge about climate change impacts on the native fauna and 

flora of grasslands, shrublands and deserts of the interior 

American West. Species’ responses will depend not only on 

their physiological tolerances but also on their phenology, 

establishment properties, biotic interactions, and capacity to 

evolve and migrate. The volume is divided into eight chapters that cover the topics of carbon mitigation and adaptation. Current and likely responses of species and habitats to 

climate change are examined in relation to taxonomic group 

and ecoregion and with regard to other disturbances. The 

volume ends with a review of management decision support needs and tools for assessing vulnerability of natural 

resources and conserving and restoring ecosystems that are 

or may be impacted by climate change.

Link: http://www.treesearch.fs.fed.us/pubs/41171

63. Franks SJ, Weis AE. 2008. A change in climate 

causes rapid evolution of multiple life-history traits and their interactions in an annual plant. Journal of Evolutionary Biology 

21:1321-1334.

Type: Journal 

Geographic Area: Southwestern USA

Keywords: Brassica rapa, common principal components analysis, developmental constraints, drought, global 

change, heterochrony, multivariate evolution, natural selection, phenology, resurrection paradigm

Abstract: Climate change is likely to spur rapid evolution, 

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 23

Keywords: climate change, forests, long-term studies, northeastern United States, winter

Abstract: Evaluations of the local effects of global change are 

often confounded by the interactions of natural and anthropogenic factors that overshadow the effects of climate changes on 

ecosystems. Long-term watershed and natural elevation gradient studies at the Hubbard Brook Experimental Forest and in 

the surrounding region show surprising results demonstrating the effects of climate change on hydrologic variables (e.g., 

evapotranspiration, streamflow, soil moisture); the importance 

of changes in phenology on water, carbon, and nitrogen fluxes 

during critical seasonal transition periods; winter climate change 

effects on plant and animal community composition and ecosystem services; and the effects of anthropogenic disturbances 

and land-use history on plant community composition. These 

studies highlight the value of long-term integrated research for 

assessments of the subtle effects of changing climate on complex ecosystems.

Link: http://www.treesearch.fs.fed.us/pubs/42303

67. Guisan A, Thuiller W. 2005. Predicting species 

distribution: offering more than simple habitat models.

Ecology Letters 8:993-1009.

Type: Journal 

Geographic Area: Global

Keywords: dispersal, ecological niche theory, future projections, habitat suitability maps, population dynamics, prediction 

errors, predictive biogeography, spatial scales, species distribution models

Abstract: In the last two decades, interest in species distribution models (SDMs) of plants and animals has grown dramatically. Recent advances in SDMs allow us to potentially forecast 

anthropogenic effects on patterns of biodiversity at different spatial scales. However, some limitations still preclude the use of 

SDMs in many theoretical and practical applications. Here, we 

provide an overview of recent advances in this field, discuss the 

ecological principles and assumptions underpinning SDMs, and 

highlight critical limitations and decisions inherent in the construction and evaluation of SDMs. Particular emphasis is given 

to the use of SDMs for the assessment of climate change impacts 

and conservation management issues. We suggest new avenues 

for incorporating species migration, population dynamics, biotic 

interactions and community ecology into SDMs at multiple spatial scales. Addressing all these issues requires a better integration of SDMs with ecological theory.

Link: http://perceval.bio.nau.edu/downloads/grail/climate_

seminar/section2/Guisan_and_Thuiller05.pdf

68. Hanson HC. 1958. Principles concerned in the formation and classification of communities. The Botanical Review 

24:65-125.

decisive in determining the success of many species, as physiological processes in C3-plants display near-linear relationships with temperatures in the range which prevails for most 

of the time. The life-cycle of native plants responds to 

environmental cues in such a way as to synchronize the 

development of the plant with the succession of seasons. 

When species are transplanted to different phytogeographical zones, as in forestry and horticulture, they frequently fail 

because of asynchrony.

Link: http://onlinelibrary.wiley.com/

doi/10.1111/j.1469-8137.1987.tb04686.x/pdf

65. Grant GE, Tague CL, Allen CD. 2013. Watering 

the forest for the trees: an emerging priority for managing 

water in forest landscapes. Frontiers in Ecology and the 

Environment 11:314-321.

Type: Journal

Geographic Area: North America

Compilers’ Keywords: forest management, hydrology, climate change, drought, water management, conservation

Abstract: Widespread threats to forests resulting from 

drought stress are prompting a re-evaluation of priorities for 

water management on forest lands. In contrast to the widely 

held view that forest management should emphasize providing water for downstream uses, we argue that maintaining forest health in the context of a changing climate may require 

focusing on the forests themselves and on strategies to reduce 

their vulnerability to increasing water stress. Management 

strategies would need to be tailored to specific landscapes but 

could include thinning, planting and selecting for droughttolerant species, irrigating, and making more water available 

to plants for transpiration. Hydrologic modeling reveals that 

specific management actions could reduce tree mortality due 

to drought stress. Adopting water conservation for vegetation as a priority for managing water on forested lands would 

represent a fundamental change in perspective and potentially 

involve trade-offs with other downstream uses of water.

Link: http://www.fsl.orst.edu/wpg/pubs/13_Granteal_WFFT.

pdf

66. Groffman PM, Rustad LE, Templer PH, 

Campbell JL, Christenson LM, Lany NK, Socci 

AM, Vadeboncoeur MA, Schaberg PG, Wilson GF, 

Driscoll CT, Fahey TJ, Fisk MC, Goodale CL, Green 

MB, Hamburg SP, Johnson CE, Mitchell MJ, Morse 

JL, Pardo LH, Rodenhouse NL. 2012. Long-term integrated studies show complex and surprising effects of climate change in the northern hardwood forest. BioScience 

62:1056-1066.

Type: Journal 

Geographic Area: Northeastern USA

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
24 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

158 annual species occurring along a gradient of rainfall variability and aridity in Israel was used to ask the following question: 

Does average seed mass and dormancy of annual populations 

and communities decrease with increasing aridity and rainfall 

unpredictability? Soil seed bank samples were collected at the 

end of the summer drought, before the onset of the rains, from 

four plant communities. Germination was tested under irrigated 

conditions during three consecutive germination seasons to determine the overall seed germinability in each soil sample. Seed 

mass was obtained from newly produced seeds collected at the 

study sites in late spring. The community level results showed 

that, in contrast to common theoretical knowledge, seed mass 

and dormancy of the dominant annual species decreased with 

increasing aridity and rainfall variability. Accordingly, a negative correlation was found between seed mass and seed germination fractions. The present study demonstrates that an analysis 

of seed traits along climatic gradients is significantly improved 

by approaches that target both population and community levels 

simultaneously. A critical evaluation sheds new light upon the 

selective pressures that act on seed ecology of annuals along a 

climatic gradient and facilitates formulation of more mechanistic hypotheses about factors governing critical seed traits.

Link: http://www.researchgate.net/publication/216466974_

Seed_mass_and_dormancy_of_annual_plant_populations_

and_communities_decreases_with_aridity_and_rainfall_predictability/file/9fcfd509bd306822a5.pdf

70. Haughian SR, Burton PJ, Taylor SW, Curry CL. 

2012. Expected effects of climate change on forest disturbance 

regimes in British Columbia. BC Journal of Ecosystems and 

Management 13:1-24.

Type: Journal 

Geographic Area: Western Canada

Keywords: disturbance ecology, ecoprovinces, forest pathogens, insect outbreaks, tree mortality, wildfire, wind damage

Abstract: In this article we summarize the changes to forest 

disturbance regimes and forest damage that are projected to 

emerge under a changing climate in British Columbia (BC). We 

focus on regionally-specific expectations so that land managers can take pro-active steps to avoid or adapt to future conditions. While some projections are based on extrapolations of 

recent multi decadal trends, most are based on global climate 

models (GCMs) that utilize a range of scenarios for possible 

atmospheric greenhouse gas emission trajectories over the next 

century. Regardless of the models or emission scenarios used, 

it is universally expected that BC will experience warmer air 

temperatures. Projections for precipitation are more variable, 

ranging from slight decreases in some regions to substantial 

increases in others, which have different effects on disturbance 

projections. Forest fires are expected to be more frequent 

and more intense in the southern half of the province and in 

the Taiga Plains, but less important in other portions of the 

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: plant ecology, population, genetics

Summary: Characteristics of species that are important in the 

formation of plant groupings are classified as basic principles, 

as follows: Every species has certain relations to the physical 

environment: (a) essential requirements, (b) range of ecological amplitude, (c) capacity to utilize available resources. Every 

plant has relations with other organisms: (a) competition, (b) association of species, (c) reproduction and dispersal, (d) changes 

in relations because of grazing, mowing or other disturbance, 

(e) parasitism, (f) mutualism and commensalism. The ecological success of a species or a population depends upon its capacity to cope with its physical environment and associated organisms in relationships, such as those stated in these principles. 

Groupings of species in stands or communities are formed 

because of similarities and differences among them with regard 

to these principles and because of similarities and differences in 

habitats. The best descriptions are based upon analysis which 

reveals intrinsic characteristics of the stands. The more detailed 

and accurate the analysis, the more valuable is the description. 

Stands that show the greatest number of characteristics in common may be grouped into community types or abstract communities of various kinds. Precise similarity of stands in each kind 

of type is not essential. Various classifications of stands and 

community-types may be made, depending upon the purpose 

to be served. Some of the bases of classification are: similarities 

and differences in composition and structure, or in complexity 

of communities; relations to habitat, succession or geographic 

distribution; physiognomic appearance; and economic values. 

Each of these is discussed with examples.

69. Harel C, Holzapfel C, Sternberg M. 2011. Seed 

mass and dormancy of annual plant populations and communities decrease with aridity and rainfall predictability. Basic and 

Applied Ecology 12:674-684.

Type: Journal 

Geographic Area: Israel, Middle East

Keywords: desert, germination strategies, Mediterranean, 

plant community, seed bank, seed size

Abstract: Several theoretical and empirical studies have examined the influence of environmental conditions on seed traits and 

germination strategies of annual species. A positive relationship 

between seed mass and dormancy has been described for annuals occupying climatically unpredictable ecosystems. Largerseeded species tend to have higher seedling survival rates, 

while dormancy allows a bet-hedging strategy in unpredictable 

environments. Until now, these ideas have been addressed primarily for only one or a few focal species, without considering 

differences among populations and communities. The novelty 

of the present study lies in the population and community-level 

approach, where a comprehensive seed trait database including 

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 25

72. Hennon PE, D’Amore DV, Schaberg PG, 

Wittwer DT, Shanley CS. 2012. Shifting climate, altered 

niche, and a dynamic conservation strategy for yellow-cedar in 

the North Pacific Coastal Rainforest. BioScience 62:147-158.

Type: Journal 

Geographic Area: Western Canada

Keywords: climate adaptation, forest decline, root cold tolerance, Callitropsis nootkatensis, Chamaecyparis nootkatensis

Abstract: The extensive mortality of yellow-cedar along 

more than 1,000 kilometers of the northern Pacific coast of 

North America serves as a leading example of climate effects on a forest tree species. In this article, we document 

our approaches to resolving the causes of tree death, which 

we explain as a cascade of interacting topographic, foreststructure, and microclimate factors that act on a unique 

vulnerability of yellow-cedar to fine-root freezing. The 

complex causes of tree mortality are reduced to two risk factors—snow depth and soil drainage—which are then used to 

model present and future cedar habitat suitability. We propose a dynamic, comprehensive conservation strategy for 

this valuable species on the basis of zones created by shifting 

climate, cedar’s ecological niche, and observed risk factors. 

Research on yellow-cedar decline is offered as a template 

for understanding and adapting to climate change for other 

climate-forest issues. 

Link: http://www.nrs.fs.fed.us/pubs/jrnl/2012/nrs_2012_hennon_001.pdf

73. Hobbs RJ, Higgs E, Harris JA. 2009. Novel ecosystems: implications for conservation and restoration. Trends in 

Ecology and Evolution 24:599-605.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: climate change, environmental monitoring, human activities, models, population density, ecosystem 

function, migration

Abstract: Many ecosystems are rapidly being transformed 

into new, non-historical configurations owing to a variety of 

local and global changes. We discuss how new systems can 

arise in the face of primarily biotic change (extinction and/or 

invasion), primarily abiotic change (e.g. land use or climate 

change) and a combination of both. Some changes will result 

in hybrid systems retaining some original characteristics as 

well as novel elements, whereas larger changes will result 

in novel systems, which comprise different species, interactions and functions. We suggest that these novel systems will 

require significant revision of conservation and restoration 

norms and practices away from the traditional place-based 

focus on existing or historical assemblages.

Link: http://www.ncbi.nlm.nih.gov/pubmed/19683830

province. Forest insects and fungal pathogens are expected to 

more fully occupy the current range of their host tree species 

and expand ranges northward and to higher elevations along 

with their hosts. More frequent and more detrimental pest outbreaks are expected in some regions when several years of favourable weather align, which is more likely under current and 

projected climate trends. Wind damage, floods, and landslides 

can be expected to increase on terrain where they are already 

a risk factor. For many agents of tree mortality, an expansion 

or shifting of the seasonal window of activity is expected, but 

these changes vary among regions within BC. The prediction 

of future forest disturbance regimes is in its infancy, requiring 

a much more concerted effort in compiling both empirical and 

simulated data, but managers may wish to adjust plans accordingly where there is consensus among projections.

Link: http://www.jem.forrex.org/index.php/jem/article/

viewArticle/152

71. Heikkinen RK, Luoto M, Araújo MB, Virkkala 

R, Thuiller W, Sykes MT. 2006. Methods and uncertainties in bioclimatic envelope modelling under climate change. 

Progress in Physical Geography 20:751-777.

Type: Journal 

Geographic Area: Global

Keywords: bioclimatic model, climate change, land cover, 

model performance, modelling methods, niche properties, 

scale, species distribution model, species geography, uncertainty, validation

Abstract: Potential impacts of projected climate change on 

biodiversity are often assessed using single-species bioclimatic 

‘envelope’ models. Such models are a special case of species 

distribution models in which the current geographical distribution of species is related to climatic variables so to enable projections of distributions under future climate change scenarios. This 

work reviews a number of critical methodological issues that 

may lead to uncertainty in predictions from bioclimatic modelling. Particular attention is paid to recent developments of bioclimatic modelling that address some of these issues as well as 

to the topics where more progress needs to be made. Developing 

and applying bioclimatic models in an informative way requires 

good understanding of a wide range of methodologies, including 

the choice of modelling technique, model validation, collinearity, autocorrelation, biased sampling of explanatory variables, 

scaling and impacts of nonclimatic factors. A key challenge for 

future research is integrating factors such as land cover, direct 

CO2 effects, biotic interactions and dispersal mechanisms into 

species-climate models. We conclude that, although bioclimatic 

envelope models have a number of important advantages, they 

need to be applied only when users of models have a thorough 

understanding of their limitations and uncertainties.

Link: http://citeseerx.ist.psu.edu/viewdoc/download?doi=10.

1.1.105.4455&rep=rep1&type=pdf

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
26 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Abstract: For tree species, adult survival and seedling and 

sapling recruitment dynamics are the main processes that 

determine forest structure and composition. Thus, studying how these two life stages may be affected by climate 

change in the context of other abiotic and biotic variables 

is critical to understand future population trends. The aim 

of this study was to assess the sustainability of cork oak 

(Quercus suber) forests at the core of its distributional 

range under future climatic conditions. Using forest inventory data collected at two periods 10 years apart, we 

performed a comprehensive analysis to evaluate the role 

of different abiotic and biotic factors on adult survival and 

recruitment patterns. We found that both life stages were 

influenced by climatic conditions, but in different ways. 

Adult tree survival was negatively impacted by warmer 

spring temperatures, while recruitment was positively affected by warmer winter temperatures. Our results also 

revealed the importance of soil texture as a modulator of 

winter precipitation effects on adult survival. With higher 

winter precipitation, adult survival increased in sandy soils 

and decreased in clayish soils. Therefore, under predicted 

future climate scenarios of wetter winters and warmer temperatures, the presence of cork oaks is more likely to occur 

in sandy soils vs. clayish soils. Biotic conditions also affected these life stages. We found a negative effect of heterospecific but not conspecific trees on both adult survival 

and seedling recruitment. Overall, the sustainability of the 

studied forests will be highly dependent not only on future 

climatic trends, but also on their interaction with other key 

factors—soil properties in particular—that modulate the 

effects of climate on demographic rates.

Link: http://www.diverbos.com/wp-content/uploads/2014/07/

Ibanez_2014_DD_wAppendix.pdf

76. Iverson LR, Matthews SN, Prasad AM, Peters 

MP, Yohe G. 2012. Development of risk matrices for evaluating climatic change responses of forested habitats. Climatic 

Change 114:231-243.

Type: Journal 

Geographic Area: Eastern USA

Compilers’ Keywords: extinction, migration, approach

Abstract: We present an approach to assess and compare risk 

from climate change among multiple species through a risk matrix, in which managers can quickly prioritize for species that 

need to have strategies developed, evaluated further, or watched. 

We base the matrix upon earlier work towards the National 

Climate Assessment for potential damage to infrastructures 

from climate change. Risk is defined here as the product of the 

likelihood of an event occurring and the consequences or impact 

of that event. In the context of species habitats, the likelihood 

component is related to the potential changes in suitable habitat modeled at various times during this century. Consequences 

74. Holmes J, Lowe J, Wolff E, Srokosz M. 2011.

Rapid climate change: lessons from the recent geological past. 

Global and Planetary Change 79:157-162.

Type: Journal 

Geographic Area: Global

Keywords: rapid climate change, Late Quaternary, thermohaline circulation, meridional overturning circulation, 8.2 ka event, 

Termination I 

Abstract: Rapid, or abrupt, climate change is regarded as 

a change in the climate system to a new state following the 

crossing of a threshold. It generally occurs at a rate exceeding 

that of the change in the underlying cause. Episodes of rapid 

climate change abound in the recent geological past (defined 

here as the interval between the last glacial maximum, dated 

to approximately 20,000 years ago, and the present). Rapid 

climate changes are known to have occurred over time periods 

equal to or even less than a human lifespan: moreover, their 

effects on the global system are sufficiently large to have had 

significant societal impacts. The potential for similar events to 

occur in the future provides an important impetus for investigating the nature and causes of rapid climate change. This 

paper provides a brief overview of rapid climate change and 

an introduction to this special issue, which presents results 

generated by the palaeoclimatic component of the UK Natural 

Environment Research Council’s rapid climate change programme, called RAPID. The papers in the special issue employ 

palaeoclimatic proxy data-sets obtained from marine, ice core 

and terrestrial archives to reconstruct rapid climate change 

during the last glacial cycle, its subsequent termination and 

the ensuing Holocene interglacial; some papers also report 

new attempts to match the palaeoclimate data to hypothesised 

causes through numerical modelling. The results confirm the 

importance of freshwater forcing in triggering changes in 

Atlantic meridional overturning circulation (MOC) and the 

close Links between MOC and rapid climate change. While 

advancing our understanding of these Linkages, the RAPID 

research has highlighted the need for further research in order 

to elucidate more specific details of the mechanisms involved.

Link: http://www.see.ed.ac.uk/~shs/Climate%20change/

Data%20sources/Rapid%20climate%20change%20NERC.pdf

75. Ibáñez B, Ibáñez I, Gómez-Aparicio L, RuizBenito P, García LV, Marañón T, Thuiller W. 2014.

Contrasting effects of climate change along life stages of a dominant tree species: the importance of soil-climate interactions.

Diversity and Distributions 20:872-883.

Type: Journal 

Geographic Area: Southern Spain

Keywords: Bayesian analysis, declining forest, demographic rates, establishment, forest inventory data, 

Mediterranean region

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 27

responses of plants grown in pots. The availability of field 

chambers for growing trees, and long-term exposure studies 

of tree species to elevated carbon dioxide, has changed much 

of our views on carbon dioxide acting as a fertiliser. Several 

tree species showed acclimation or even down-regulation of 

photosynthetic responses while a few of them showed higher 

photosynthesis and better growth responses. Whether elevated 

levels of carbon dioxide can serve as a fertilizer in a changed 

climate scenario still remains an unresolved question. ForestAir-Carbon dioxide Enrichment (FACE) sites monitored at 

several locations have shown lately, that the acclimation or 

down regulation as reported in chamber studies is not as widespread as originally thought. FACE studies predict that there 

could be an increase of 23-28% productivity of trees at least 

till 2050. However, the increase in global temperature could 

also lead to increased respiration, and limitation of minerals in 

the soil could lead to reduced responses in growth. Elevated 

carbon dioxide induces partial closure of leaf stomata, which 

could lead to reduced transpiration and more economical use 

of water by the trees. Even if the carbon dioxide acts as a fertilizer, the responses are more pronounced only in young trees. 

And if there are variations in species responses to growth due 

to elevated carbon dioxide, only some species are going to 

dominate the natural vegetation. This will have serious implications on the biodiversity and the structure of the ecosystems. 

This paper reviews the research done on trees using elevated 

CO2 and tries to draw conclusions based on different methods 

used for the study. It also discusses the possible functional 

variations in some tree species due to climate change. 

79. Keenan TF, Hollinger DY, Bohrer G, Dragoni 

D, Munger JW, Schmid HP, Richardson AD. 2013.

Increase in forest water-use efficiency as atmospheric carbon dioxide concentrations rise. Nature 499:324-327.

Type: Journal 

Geographic Area: Northern Hemisphere

Compilers’ Keywords: ecosystem, plant leaves, trees

Abstract: Terrestrial plants remove CO2 from the atmosphere 

through photosynthesis, a process that is accompanied by the 

loss of water vapour from leaves. The ratio of water loss to 

carbon gain, or water-use efficiency, is a key characteristic of 

ecosystem function that is central to the global cycles of water, 

energy and carbon. Here we analyse direct, long-term measurements of whole-ecosystem carbon and water exchange. 

We find a substantial increase in water-use efficiency in temperate and boreal forests of the Northern Hemisphere over the 

past two decades. We systematically assess various competing hypotheses to explain this trend, and find that the observed 

increase is most consistent with a strong CO2 fertilization 

effect. The results suggest a partial closure of stomata-small 

pores on the leaf surface that regulate gas exchange-to maintain a near-constant concentration of CO2 inside the leaf even 

under continually increasing atmospheric CO2 levels. The 

are related to the adaptability of the species to cope with the 

changes, especially the increasing intensity and/or frequency of 

disturbance events that are projected. We derived consequence 

scores from nine biological and 12 disturbance characteristics 

that were pulled from literature for each species. All data were 

generated from an atlas of climate change for 134 trees of the 

eastern United States. We show examples which depict a wide 

range of risk for tree species of northern Wisconsin, including 

species that may gain substantial habitat as well as lose substantial habitat, both of which will require the development of strategies to help the ecosystems adapt to such changes. 

Link: http://www.nrs.fs.fed.us/pubs/jrnl/2012/nrs_2012_iverson_001.pdf

77. Jones EA, Reed DD, Desanker PV. 1994. Ecological 

implications of projected climate change scenarios in forest 

ecosystems of central North America. Agricultural and Forest 

Meteorology 72:31-46.

Type: Journal 

Geographic Area: Central North America

Compilers’ Keywords: climate change impacts, projections, 

commercial tree species, GCM, weather stations

Abstract: Climate change scenarios in central North America 

were projected for selected weather stations using a stochastic 

daily weather simulation model. The projections were conditioned on changes in seasonal temperature and precipitation 

as predicted by several global climate models. The impacts of 

these projected changes on temperate forest ecosystems were 

evaluated through projected changes in such variables as average annual temperature, average growing season temperature, 

and the ratio of potential evaporation to precipitation during July 

and August. Even the mildest climate change scenario examined 

indicated that ecologically significant changes could occur in 

the composition and productivity of these forests. The possibility also exists that climatically induced regional decline episodes 

for a number of important commercial species could occur in the 

northern temperate forests of central North America.

Link: http://www.webpages.uidaho.edu/nitrogen-gradient/

Publications/Jones_1994_Agric_For_Metogy.pdf

78. Kallarackal J, Roby TJ. 2012. Responses of trees to 

elevated carbon dioxide and climate change. Biodiversity and 

Conservation 21:1327-1342.

Type: Journal 

Geographic Area: Global

Keywords: trees, climate change, responses, carbon dioxide, 

temperature, growth, FACE

Abstract: The enhancement in photosynthesis at elevated 

concentration of carbon dioxide level than the ambient level existing in the atmosphere is widely known. However, 

many of the earlier studies were based on the instantaneous 

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
28 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Link: http://research.eeescience.utoledo.edu/lees/papers_pdf/

Kramer_2000_IntJBiomet.pdf

81. Kreyling J. 2010. Winter climate change: a critical factor 

for temperate vegetation performance. Ecology 97:1939-1948.

Type: Journal 

Geographic Area: Global

Keywords: climate change, frost, global warming, snow, temperate zone, winter ecology

Abstract: Winter ecological processes are important drivers of 

vegetation and ecosystem functioning in temperate ecosystems. 

There, winter conditions are subject to rapid climate change. 

The potential loss of a longer-lasting snow cover with implications to other plant-related climate parameters and overwintering strategies make the temperate zone particularly vulnerable 

to winter climate change. A formalized literature search in the 

ISI Web of Science shows that plant related research on the 

effects of winter climate change is generally underrepresented. 

Temperate regions in particular are rarely studied in this respect, although the few existing studies imply strong effects 

of winter climate change on species ranges, species compositions, phenology, or frost injury. The generally positive effect 

of warming on plant survival and production may be counteracted by effects such as an increased frost injury of roots and 

shoots, an increased insect pest risk, or a disrupted synchrony 

between plants and pollinators. Based on the literature study, 

gaps in current knowledge are discussed. Understanding the 

relative effects of interacting climate parameters, as well as a 

stronger consideration of short-term events and variability of 

climatic conditions is urgent. With respect to plant response, it 

would be particularly worthwhile to account for hidden players 

such as pathogens, pollinators, herbivores, or fungal partners 

in mycorrhization.

Link: ftp://norbif.uio.no/pub/outgoing/runeho/KR/

Krey10Ecol91-1939.pdf

82. Li J, Wang MH, Ho YS. 2011. Trends in research on 

global climate change: a Science Citation Index Expanded-based 

analysis. Global and Planetary Change 77:13-20.

Type: Journal 

Geographic Area: Global

Keywords: word cluster analysis, bibliometrics, climate 

change, research trend, model

Abstract: This study was conceived to evaluate the global 

scientific output of climate change research over the past 18 

years and to assess the characteristics of the research patterns, 

tendencies, and methods in the papers. Data were based on the 

online version of Science Citation Index Expanded from 1992 

to 2009. Articles referring to climate change were assessed by 

distribution of source countries, source institutes, paper titles, 

author keywords, KeyWords Plus, abstracts, and the most cited 

observed increase in forest water-use efficiency is larger than 

that predicted by existing theory and 13 terrestrial biosphere 

models. The increase is associated with trends of increasing 

ecosystem-level photosynthesis and net carbon uptake, and decreasing evapotranspiration. Our findings suggest a shift in the 

carbon— and water—based economics of terrestrial vegetation, which may require a reassessment of the role of stomatal 

control in regulating interactions between forests and climate 

change, and a re-evaluation of coupled vegetation-climate 

models.

Link: http://www.ncbi.nlm.nih.gov/pubmed/23842499

80. Kramer K, Leinonen I, Loustau D. 2000.

The importance of phenology for the evaluation of impact of climate change on growth of boreal, temperate and Mediterranean 

forests ecosystems: an overview. International Journal of 

Biometeorology 44:67-75.

Type: Journal 

Geographic Area: Europe

Keywords: annual cycle, climate change scenarios, frost hardiness, growth, forest growth model, photosynthetic capacity, 

competition, water availability

Abstract: An overview is presented of the phenological models relevant for boreal coniferous, temperate-zone deciduous 

and Mediterranean coniferous forest ecosystems. The phenology of the boreal forests is mainly driven by temperature, affecting the timing of the start of the growing season and thereby its duration, and the level of frost hardiness and thereby the 

reduction of foliage area and photosynthetic capacity by severe 

frost events. The phenology of temperate-zone forests is also 

mainly driven by temperature. Since temperate-zone forests are 

mostly mixed-species deciduous forests, differences in phenological response may affect competition between tree species. 

The phenology of Mediterranean coniferous forests is mainly 

driven by water availability, affecting the development of leaf 

area, rather than the timing of phenological events. These phenological models were subsequently coupled to the processbased forest model FORGRO to evaluate the effect of different 

climate change scenarios on growth. The results indicate that 

the phenology of each of the forest types significantly affects 

the growth response to a given climate change scenario. The 

absolute responses presented in this study should, however, be 

used with caution as there are still uncertainties in the phenological models, the growth models, the parameter values obtained and the climate change scenarios used. Future research 

should attempt to reduce these uncertainties. It is recommended that phenological models that describe the mechanisms by 

which seasonality in climatic drivers affects the phenological 

aspects of trees should be developed and carefully tested. Only 

by using such models may we make an assessment of the impact of climate change on the functioning and productivity of 

different forest ecosystems.

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 29

also be impacted by climate change, but much less knowledge is available to quantify these impacts. Understanding of 

adaptive capacity and regional vulnerability to climate change 

in European forests is not well developed and requires more 

focused research efforts. An interdisciplinary research agenda 

integrated with monitoring networks and projection models is 

needed to provide information at all levels of decision making, 

from policy development to the management unit.

Link: http://citeseerx.ist.psu.edu/viewdoc/download?doi=10.

1.1.466.7250&rep=rep1&type=pdf

84. Littell J, Peterson DL, Tjoelker M. 2008.Douglasfir growth in mountain ecosystems: water limits tree growth 

from stand to region. Ecological Monographs 78:349-368.

Type: Journal 

Geographic Area: Northwestern USA

Keywords: climate change, climate effects, dendrochronology, Douglas-fir, ecological amplitude, ecophysiology, limiting factors, niche, Pseudotsuga menziesii, tree growth, water 

balance deficit

Abstract: The purpose of this work is to understand the nature 

of growth–climate relationships for Douglas-fir (Pseudotsuga 

menziesii) across the climatic dimensions of its niche. We used 

a combination of biophysically informed sampling (to identify sample sites) and dendroclimatology (to identify growth–

climate relationships) along a climate gradient in northwestern 

United States mountain ecosystems from the western Olympic 

Peninsula, Washington to the eastern Rocky Mountain Front, 

Montana. We used a multi-scale sampling strategy that accounted for continentality, physiography, and topography as 

non-climatic factors that could influence climate and alter tree 

growth. We developed a network of 124 Douglas-fir tree-ring 

chronologies and explored growth–climate correlations across 

the sampled gradients. We considered two different spatial 

scales of monthly and seasonal climate variables as potential 

controlling factors on tree growth. Annual radial growth in 

60–65% of the plots across the region is significantly correlated with precipitation, drought, or water balance during the 

late summer prior to growth and the early summer the year 

of growth. In a few plots, growth is positively correlated with 

cool-season temperature or negatively correlated with snowpack. Water availability is therefore more commonly limiting to Douglas-fir growth than energy limitations on growing 

season length. The first principal component derived from 

the chronologies is significantly correlated with independent 

drought reconstructions. The sensitivity of Douglas-fir to summer water balance deficit (potential evapotranspiration minus 

actual evapotranspiration) indicates that increases in April to 

September temperature without increases in summer precipitation or soil moisture reserves are likely to cause decreases 

in growth over much of the sampled area, especially east of 

the Cascade crest. In contrast, Douglas-fir may exhibit growth 

articles in these years. By synthetic analysis of the four kinds 

of keywords, it was concluded that the items “temperature,” 

“environment,” “precipitation,” “greenhouse gas,” “risk,” and 

“biodiversity” will be the foci of climate change research in 

the 21st century, while “model,” “monitoring,” and “remote 

sensing” will continue to be the leading research methods. A 

novel method, “phylogeography,” may have a strong application potential in the near future.

Link: http://trend.asia.edu.tw/Visitors/JF%20Li/Glo%20

Pla%20Cha77,%2013.pdf

83. Lindner M, Maroschek M, Netherer S, Kremer 

A, Barbati A, Garcia-Gonzalo J, Seidl R, Delzon S, 

Corona P, Kolström M, Lexer MJ, Marchetti M. 

2010. Climate change impacts, adaptive capacity, and vulnerability of European forest ecosystems. Forest Ecology and 

Management 259:698-709.

Type: Journal 

Geographic Area: Europe

Keywords: climate change impacts, adaptive capacity, forestry, vulnerability, regional differences

Abstract: This study compiles and summarizes the existing 

knowledge about observed and projected impacts of climate 

change on forests in Europe. Forests will have to adapt not 

only to changes in mean climate variables but also to increased 

variability with greater risk of extreme weather events, such 

as prolonged drought, storms and floods. Sensitivity, potential 

impacts, adaptive capacity, and vulnerability to climate change 

are reviewed for European forests. The most important potential impacts of climate change on forest goods and services are 

summarized for the Boreal, Temperate Oceanic, Temperate 

Continental, Mediterranean, and mountainous regions. 

Especially in northern and western Europe the increasing atmospheric CO2 content and warmer temperatures are expected 

to result in positive effects on forest growth and wood production, at least in the short–medium term. On the other hand, 

increasing drought and disturbance risks will cause adverse 

effects. These negative impacts are very likely to outweigh 

positive trends in southern and eastern Europe. From west to 

east, the drought risk increases. In the Mediterranean regions 

productivity is expected to decline due to strongly increased 

droughts and fire risks. Adaptive capacity consists of the inherent adaptive capacity of trees and forest ecosystems and of socio-economic factors determining the capability to implement 

planned adaptation. The adaptive capacity in the forest sector 

is relatively large in the Boreal and the Temperate Oceanic 

regions, more constrained by socio-economic factors in the 

Temperate Continental, and most limited in the Mediterranean 

region where large forest areas are only extensively managed 

or unmanaged. Potential impacts and risks are best studied and 

understood with respect to wood production. It is clear that all 

other goods and services provided by European forests will 

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
30 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

86. Lloret F, Peñuelas J, Prieto P, Llorens L, 

Estiarte M. 2009. Plant community changes induced by experimental climate change: seedling and adult species composition. Perspectives in Plant Ecology, Evolution and Systematics 

11:53-63.

Type: Journal 

Geographic Area: Mediterranean

Keywords: climate change, drought, Mediterranean, seedling 

establishment, vegetation dynamics, warming

Abstract: Experimental manipulation of climate provides a 

powerful tool for studying plant community dynamics with 

respect to current climate change. We experimentally investigated the vegetation dynamics of a Mediterranean shrubland 

under directional climate change by manipulating rain and 

temperature at stand level throughout 7 years. We focused on 

seedling establishment in relation to the between-year variability of drought conditions. We also compared seedling dynamics to changes in the established adult vegetation to assess 

the coupling between both dynamics. We used multivariate 

techniques (principal response curves (PRC) and redundancy 

analysis (RDA)) to explore changes in the whole community, 

and Generalized Linear Model (GLZM) to analyze the influence of drought on the abundance and survival of the most 

abundant species. Drought treatment induced significant 

changes in the species composition of the seedlings, via a 

differential decrease in the seedling density of most species. 

No species was particularly favoured in terms of seedling 

abundance under water-deficit conditions. Warming only explained a low percentage of the variability in seedling species 

composition. The emergence of seedlings in control plots—

which may be considered an estimation of the between-year 

variability in the conditions for seedling establishment—was 

a better predictor of seedling emergence in experimental plots 

than climate manipulation treatments. The PRC analysis of 

the adults showed dynamics that were different from those 

recorded for seedlings, and it also showed that drought treatment significantly explained species composition. This result 

is reinforced by the change in the relative abundance of seedling and adults of the more common species in the drought and 

warming treatments, supporting the hypothesis that climatic 

directional change heightens discrepancies between recruitment and the adult performance. The RDA analysis applied 

to species composition at the end of the experiment failed, 

however, to attain any statistical significance. The warming 

treatment did not produce any significant shifts in adult vegetation. In conclusion, directional climate change—particularly drier conditions in Mediterranean shrublands—would 

result in a change in the recruitment of the plant community. 

This change in seedling recruitment tends to be different from 

the dynamics of adults, suggesting that potential adult mortality would not be compensated by actual seedling recruitment, 

thus enhancing shifts in community composition. 

increases at some higher elevation sites where seasonal photosynthesis is currently limited by growing-season length or low 

growing-season temperature. Life-history processes such as 

establishment, growth, and mortality are precursors to changes 

in biogeography, and measurements of climate effects on those 

processes can provide early indications of climate-change effects on ecosystems.

Link: http://www.fs.fed.us/pnw/pubs/journals/pnw_2008_littell001.pdf

85. Littell J, Elsner MM, Mauger GS, Lutz E, 

Hamlet AF, Salathe E. 2011. Regional climate and 

hydrologic change in the Northern U.S. Rockies and Pacific 

Northwest: internally consistent projections of future climate for 

resource management. Seattle (WA): Climate Impacts Group, 

University of Washington. 109 p.

Type: Report

Geographic Area: Western USA

Compilers’ Keywords: planning, natural resources, global 

climate model GCM, watershed

Rationale and Objectives: Planning for the effects of climate change on natural resources often requires detailed projections of future climate at scales consistent with the processes 

managers typically consider. While it is numerically possible 

to produce downscaled climate at very fine scales (< 5 km), 

both the absence of a sufficiently dense network of long term 

climate observations and the presence of local contingencies 

such as topography and land surface feedbacks from vegetation 

and snowpack make accurate estimation at these scales difficult and less tractable without very detailed local information. 

For such purposes as developing adaptation strategies, vulnerability assessments, climate impacts assessments, and specific 

resource modeling at landscape scales, downscaled projections can be developed that maximize translation of climatic 

information from the coarser scales of global climate models 

(GCMs) to more local scales. This project was designed to provide climate information that meets those needs and creates 

a basis for more detailed work or for a more comprehensive 

approach to downscaling and regional climate modeling. The 

goal of this project was to (1) develop consistent historical and 

future downscaled climate and hydrologic data and projections 

using the same methodology for several major river basins in 

the western United States (Columbia, upper Missouri, upper 

Colorado, and Great Basins) and (2) summarize that information in forms consistent with the needs of the funding agencies. 

This report describes where to get the information developed 

as well as the methods, results obtained, uses of and uncertainties associated with the data and projections. 

Link: http://cses.washington.edu/picea/USFS/pub/

Littell_etal_2010/Littell_etal._2011_Regional_Climatic_And_

Hydrologic_Change_USFS_USFWS_JVA_17Apr11_reduced.

pdf

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 31

rather than soil drought. Vulnerability to xylem cavitation did 

not differ between desert and montane populations. A 50% loss 

in hydraulic conductivity occurred at water potentials between 

–2.61 and –2.65 MPa, and vulnerability to xylem cavitation 

did not vary with stem size. Minimum xylem tensions of desert and montane trees did not drop below –2.05 MPa. Foliage 

turgor loss point did not differ between climate groups and corresponded to mean minimum xylem tensions in the field. In 

addition to low AL/AS, high KS in desert trees may provide a 

way to increase tree hydraulic conductivity in response to high 

evaporative demand and prevent xylem tensions from reaching 

values that cause catastrophic cavitation. In ponderosa pine, 

the flexible responses of AL/AS and KS to climate may preclude the existence of significant intraspecific variation in the 

vulnerability of xylem to cavitation.

Link: http://classes.uleth.ca/201301/biol3460a/Lab%20

Materials/Lab%20Presentation%20Project/Project%202/

Maherali%20TreePhysiol%202000.pdf

89. Maherali H, Williams BL, Paige KN, DeLucia 

EH. 2002. Hydraulic differentiation of ponderosa pine populations along a climate gradient is not associated with ecotypic 

divergence. Functional Ecology 16:510-521.

Type: Journal 

Geographic Area: Southwestern USA

Keywords: ecotypes, genetic variation, hydraulic architecture, microsatellites, phenotypic plasticity

Summary: Pinus ponderosa occurs in a range of contrasting 

environments in the western USA. Xeric populations typically have lower leaf : sapwood area ratio (AL/AS) and higher 

whole-tree leaf specific hydraulic conductance (KL) than mesic 

populations. These climate driven shifts in hydraulic architecture are considered adaptive because they maintain minimum 

leaf water potential above levels that cause xylem cavitation. 

Using a common garden study, we examined whether differences in biomass allocation and hydraulic architecture between 

P. ponderosa populations originating from isolated outcrops 

in the Great Basin desert and Sierran montane environments 

were caused by ecotypic differentiation or phenotypic plasticity. To determine if populations were genetically differentiated and if phenotypic and genetic differentiation coincided, 

we also characterized the genetic structure of these populations using DNA microsatellites. Phenotypic differentiation 

in growth, biomass allocation and hydraulic architecture was 

variable among populations in the common garden. There 

were no systematic differences between desert and montane 

climate groups that were consistent with adaptive expectations. Drought had no effect on the root : shoot and needle : 

stem ratio, but reduced seedling biomass accumulation, leaf 

area ratio, AL/AS and KL. Stem hydraulic conductance (KH) 

was strongly size dependent, and was lower in drought plants, 

primarily because of lower growth. Although microsatellites 

87. Logan JA, Regniere J, Powell JA. 2003. Assessing 

the impacts of global warming on forest pest dynamics. Frontiers 

in Ecology and the Environment 1:130-137.

Type: Journal 

Geographic Area: North America

Compilers’ Keywords: climate change, insect outbreaks, 

forest management, review

Abstract: Forest insects and pathogens are the most pervasive 

and important agents of disturbance in North American forests, 

affecting an area almost 50 times larger than fire and with an 

economic impact nearly five times as great. The same attributes that result in an insect herbivore being termed a “pest” 

predispose it to disruption by climate change, particularly 

global warming. Although many pest species have co-evolved 

relationships with forest hosts that may or may not be harmful over the long term, the effects on these relationships may 

have disastrous consequences. We consider both the data and 

models necessary to evaluate the impacts of climate change, 

as well as the assessments that have been made to date. The 

results indicate that all aspects of insect outbreak behavior will 

intensify as the climate warms. This reinforces the need for 

more detailed monitoring and evaluations as climatic events 

unfold. Luckily, we are well placed to make rapid progress, 

using software tools, databases, and the models that are already 

available.

Link: http://www.usu.edu/beetle/documents/Loganet.al.2003.

pdf

88. Maherali H, DeLucia EH. 2000. Xylem conductivity and vulnerability to cavitation of ponderosa pine growing in 

contrasting climates. Tree Physiology 20:859-867.

Type: Journal 

Geographic Area: Southwestern USA

Keywords: hydraulic conductivity, leaf/sapwood area ratio, 

temperature, vapor pressure deficit, xylem cavitation

Summary: We examined the effects of increased transpiration 

demand on xylem hydraulic conductivity and vulnerability to 

cavitation of mature ponderosa pine (Pinus ponderosa Laws.) 

by comparing trees growing in contrasting climates. Previous 

studies determined that trees growing in warm and dry sites 

(desert) had half the leaf/sapwood area ratio (AL/AS) and 

more than twice the transpiration rate of trees growing in cool 

and moist sites (montane). We predicted that high transpiration 

rates would be associated with increased specific hydraulic 

conductivity (KS) and increased resistance to xylem cavitation. 

Desert trees had 19% higher KS than montane trees, primarily because of larger tracheid lumen diameters. Predawn water 

potential and water potential differences between the soil and 

the shoot were similar for desert and montane trees, suggesting 

that differences in tracheid anatomy, and therefore KS, were 

caused primarily by temperature and evaporative demand, 

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
32 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

episodes of drought-induced forest dieback have been reported 

from around the world, which points to an emerging need for 

multiscale monitoring approaches to quantify drought effects 

on woody biomass and carbon cycling across large areas.

Link: http://nofc.cfs.nrcan.gc.ca/bookstore_pdfs/31947.pdf

91. Millar CI, Woolfenden WB. 1999. Sierra Nevada 

forests: where did they come from? Where are they going? What 

does it mean? Transactions of the 64th North American Wildlife 

and Natural Resources Conference. p 206-236.

Type: Conference Paper

Geographic Area: Sierra Nevada, California, North America

Compilers’ Keywords: Quaternary sciences, climate 

change, historic natural variability, vegetation distribution, 

land management

Introduction: The prospect of human-induced global warming has received center-stage attention from ecologists concerned about future ecosystems. While such effects deserve 

critical analysis, they are best understood in the context of 

natural climate change. Anthropogenic atmospheric effects are 

often discussed as if background climates were stable. Even in 

the recent past, however, climates have changed at similar rates 

and magnitudes to predicted anthropogenic changes, catalyzing significant natural changes in terrestrial ecosystems. With 

or without the complication of human effects, implications of 

climate change to conservation and management planning are 

great. Over the past twenty years, advances in Quaternary sciences have significantly improved our understanding of historic climate and its influence on biota and planetary systems. 

Although resource ecologists increasingly engage this research, much of relevance remains to be incorporated into the 

thinking of forest science and management. In this paper, we 

briefly review the nature of historic climate change, summarize conclusions relevant to ecosystems of the Sierra Nevada, 

California, and suggest implications for managing Sierran forest ecosystems. 

Link: http://www.treesearch.fs.fed.us/pubs/24289

92. Millar CI, King JC, Westfall RD, Alden HA, 

Delany DL. 2006. Late Holocene forest dynamics, 

volcanism, and climate change at Whitewing Mountain and 

San Joaquin Ridge, Mono County, Sierra Nevada, CA, USA.

Quaternary Research 66:273-287.

Type: Journal 

Geographic Area: California, USA 

Keywords: paleoecology, medieval climate, Late Holocene, 

climate change, Long Valley volcanism, Inyo Craters, forest 

history, palaeoclimatic modeling, tree-ring dating

Abstract: Deadwood tree stems scattered above treeline on 

tephra covered slopes of Whitewing Mtn (3051 m) and San 

Joaquin Ridge (3122 m) show evidence of being killed in an 

were able to detect significant non-zero (P < 0.001) levels of 

differentiation between populations, these differences were 

small and were not correlated with geographic separation or 

climate group. Estimates of genetic differentiation among populations were low (< 5%), and almost all the genetic variation 

(> 95%) resided within populations, suggesting that gene flow 

was the dominant factor shaping genetic structure. These results indicate that biomass allocation and hydraulic differences 

between desert and montane populations are not the result of 

ecotypic differentiation. Significant drought effects on leaf : 

sapwood allocation and KL suggest that phenotypic differentiation between desert and montane climates could be the result 

of phenotypic plasticity.

Link: http://www.life.illinois.edu/delucia/FEC510.pdf

90. Michaelian M, Hogg EH, Hall RJ, Arsenault 

E. 2011. Massive mortality of aspen following severe drought 

along the southern edge of the Canadian boreal forest. Global 

Change Biology 17:2084-2094.

Type: Journal 

Geographic Area: Canada

Keywords: aspen, boreal forest, climate change, dieback, 

drought, mortality, Populus tremuloides

Abstract: Drought-induced, regional-scale dieback of forests 

has emerged as a global concern that is expected to escalate under model projections of climate change. Since 2000, drought 

of unusual severity, extent, and duration has affected large areas 

of western North America, leading to regional scale dieback of 

forests in the southwestern US. We report on drought impacts 

on forests in a region farther north, encompassing the transition 

between boreal forest and prairie in western Canada. A central 

question is the significance of drought as an agent of large-scale 

tree mortality and its potential future impact on carbon cycling 

in this cold region. We used a combination of plot-based, meteorological, and remote sensing measures to map and quantify 

aboveground, dead biomass of trembling aspen (Populus tremuloides Michx.) across an 11.5 Mha survey area where drought 

was exceptionally severe during 2001–2002. Within this area, 

a satellite-based land cover map showed that aspen-dominated 

broadleaf forests occupied 2.3 Mha. Aerial surveys revealed 

extensive patches of severe mortality (> 55%) resembling the 

impacts of fire. Dead aboveground biomass was estimated at 

45 Mt, representing 20% of the total aboveground biomass, 

based on a spatial interpolation of plot-based measurements. 

Spatial variation in percentage dead biomass showed a moderately strong correlation with drought severity. In the prairielike, southern half of the study area where the drought was most 

severe, 35% of aspen biomass was dead, compared with an 

estimated 7% dead biomass in the absence of drought. Drought 

led to an estimated 29 Mt increase in dead biomass across the 

survey area, corresponding to 14 Mt of potential future carbon 

emissions following decomposition. Many recent, comparable 

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 33

Link: http://www.ncbi.nlm.nih.gov/pubmed/22596094

94. Moritz C, Agudo R. 2013. The future of species under climate change: resilience or decline? Science 341:504-508.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: fossil record, range shifts, climate 

change

Abstract: As climates change across already stressed ecosystems, there is no doubt that species will be affected, but to what 

extent and which will be most vulnerable remain uncertain. 

The fossil record suggests that most species persisted through 

past climate change, whereas forecasts of future impacts predict large-scale range reduction and extinction. Many species 

have altered range limits and phenotypes through 20th-century 

climate change, but responses are highly variable. The proximate causes of species decline relative to resilience remain 

largely obscure; however, recent examples of climate-associated species decline can help guide current management in 

parallel with ongoing research.

Link: http://www.ncbi.nlm.nih.gov/pubmed/23908228

95. Myers P, Lundrigan BL, Hoffman SMG, 

Haraminac AP, Seto SH. 2009. Climate-induced changes 

in the small mammal communities of the northern Great Lakes 

Region. Global Change Biology 15:1434-1454.

Type: Journal 

Geographic Area: Northern Great Lakes, USA

Keywords: climatic warming, Didelphis virginiana, 

Glaucomys sabrinus, Glaucomys volans, Great Lakes, Michigan, 

Peromyscus leucopus, Peromyscus maniculatus gracilis, Tamias 

minimus, Tamias striatus

Abstract: We use museum and other collection records to 

document large and extraordinarily rapid changes in the 

ranges and relative abundance of nine species of mammals 

in the northern Great Lakes region (white footed mice, woodland deer mice, southern red-backed voles, woodland jumping mice, eastern chipmunks, least chipmunks, southern flying squirrels, northern flying squirrels, common opossums). 

These species reach either the southern or the northern limit 

of their distributions in this region. Changes consistently reflect increases in species of primarily southern distribution 

(white-footed mice, eastern chipmunks, southern flying squirrels, common opossums) and declines by northern species 

(woodland deer mice, southern red-backed voles, woodland 

jumping mice, least chipmunks, northern flying squirrels). 

White-footed mice and southern flying squirrels have extended their ranges over 225 km since 1980, and at particularly 

well-studied sites in Michigan’s Upper Peninsula, small mammal assemblages have shifted from numerical domination by 

eruption from adjacent Glass Creek Vent, Inyo Craters. Using 

tree-ring methods, we dated deadwood to AD 815–1350 and 

infer from death dates that the eruption occurred in late summer 

AD 1350. Based on wood anatomy, we identified deadwood species as Pinus albicaulis, P. monticola, P. lambertiana, P. contorta, P. 

jeffreyi, and Tsuga mertensiana. Only P. albicaulis grows at these 

elevations currently; P. lambertiana is not locally native. Using 

contemporary distributions of the species, we modeled paleoclimate during the time of sympatry to be significantly warmer 

(+3.2 °C annual minimum temperature) and slightly drier (−24 

mm annual precipitation) than present, resembling values projected for California in the next 70–100 yr.

Link: http://www.treesearch.fs.fed.us/pubs/31776

93. Mondoni A, Rossi G, Orsenigo S, Probert RJ. 

2012. Climate warming could shift the timing of seed germination in alpine plants. Annals of Botany 110:155-164.

Type: Journal 

Geographic Area: Italy, Europe

Keywords: seed ecology, seed germination, climate change, 

adaptation, alpine plants

Abstract: Despite the considerable number of studies on the 

impacts of climate change on alpine plants, there have been few 

attempts to investigate its effect on regeneration. Recruitment 

from seeds is a key event in the life-history of plants, affecting their spread and evolution and seasonal changes in climate 

will inevitably affect recruitment success. Here, an investigation was made of how climate change will affect the timing 

and the level of germination in eight alpine species of the glacier foreland. Using a novel approach which considered the 

altitudinal variation of temperature as a surrogate for future 

climate scenarios, seeds were exposed to 12 different cycles 

of simulated seasonal temperatures in the laboratory, derived 

from measurements at the soil surface at the study site. Under 

present climatic conditions, germination occurred in spring, 

in all but one species, after seeds had experienced autumn 

and winter seasons. However, autumn warming resulted in a 

significant increase in germination in all but two species. In 

contrast, seed germination was less sensitive to changes in 

spring and/or winter temperatures, which affected only three 

species. Climate warming will lead to a shift from spring to 

autumn emergence but the extent of this change across species 

will be driven by seed dormancy status. Ungerminated seeds 

at the end of autumn will be exposed to shorter winter seasons 

and lower spring temperatures in a future, warmer climate, but 

these changes will only have a minor impact on germination. 

The extent to which climate change will be detrimental to regeneration from seed is less likely to be due to a significant 

negative effect on germination per se, but rather to seedling 

emergence in seasons that the species are not adapted to experience. Emergence in autumn could have major implications 

for species currently adapted to emerge in spring.

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
34 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Abstract: Understanding how vulnerable forest ecosystems 

are to climate change is a key requirement if sustainable forest management is to be achieved. Modelling the response of 

species in their regeneration niche to phenological and biophysical processes that are directly influenced by climate is 

one method for achieving this understanding. A model was 

developed to investigate species resilience and vulnerability to 

climate change within its fundamental regeneration niche. The 

utility of the developed model, tree and climate assessment 

(TACA), was tested within the interior Douglas-fir ecosystem 

in south central British Columbia. TACA modelled the current potential tree species composition of the ecosystem with 

high accuracy and modelled significant responses amongst tree 

species to climate change. The response of individual species 

suggests that the studied ecosystem could transition to a new 

ecosystem over the next 100 years. TACA showed that it can 

be an effective tool for identifying species resilience and vulnerability to changes in climate within the most sensitive stage 

of development, the regeneration phase. The TACA model was 

able to identify the degree of change in phenological and biophysical variables that control tree establishment, growth and 

persistence. The response to changes in one or more of these 

variables resulted in changes in the climatic suitability of the 

ecosystem for species and enabled a measure of vulnerability 

to be quantified. TACA could be useful to forest managers as a 

decision support tool for adaptation actions and by researchers 

interested in modelling stand dynamics under climate change.

Link: http://www4.nau.edu/direnet/publications/

publications_n/files/Nitschke_CR_Innes_JL_A_tree_and_climate_assessment_tool.pdf

98. Niven DK, Butcher GS, Bancroft GT, Monahan 

WB, Langham G. 2009. Birds and climate change: ecological disruption in motion. New York (NY): National Audubon 

Society. 16 p.

Type: Report

Geographic Area: North America

Compilers’ Keywords: wildlife, Audubon Society, biological indicators

Introduction: Each year for more than a century, dedicated 

volunteers have braved snow, wind, rain and ice to record the 

number and location of North American birds. The carefully 

organized and compiled observations of tens of thousands of 

Citizen Scientists participating in Audubon’s annual Christmas 

Bird Count have grown to form the world’s longest uninterrupted repository of bird population information. Analyses of 

its data have time after time revealed important trends, alerting America to perils and opportunities with implications far 

beyond avian well-being. Birds are well-known barometers 

of environmental health. Changes in their condition can warn 

of threats to habitats and natural systems critical to all life on 

Earth. Like canaries in a coal mine, they can alert us to danger. 

northern species to domination by southern species. Repeated 

resampling at some sites suggests that southern species are 

replacing northern ones rather than simply being added to the 

fauna. Observed changes are consistent with predictions from 

climatic warming but not with predictions based on recovery 

from logging or changes in human populations. Because of 

the abundance of these focal species (the eight rodent species make up 96.5% of capture records of all forest-dwelling 

rodents in the region and 70% of capture records of all forestdwelling small mammals) and the dominating ecological roles 

they play, these changes substantially affect the composition 

and structure of forest communities. They also provide an unusually clear example of change that is likely to be the result 

of climatic warming in communities that are experienced by 

large numbers of people.

Link: http://www.fws.gov/uploadedFiles/Myers2009.pdf

96. Nicotra AB, Atkin OK, Bonser SP, Davidson AM, 

Finnegan EJ, Mathesius U, Poot P, Purugganan 

MD, Richards CL, Valladares F, van Kleunen M. 

2010. Plant phenotypic plasticity in a changing climate. Trends 

in Plant Science 15:684-692.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: adaptation, climate change, flowers, 

plant physiological phenomena, plant genetics, seed physiology, review, tool

Abstract: Climate change is altering the availability of resources and the conditions that are crucial to plant performance. 

One way plants will respond to these changes is through environmentally induced shifts in phenotype (phenotypic plasticity). Understanding plastic responses is crucial for predicting 

and managing the effects of climate change on native species 

as well as crop plants. Here, we provide a toolbox with definitions of key theoretical elements and a synthesis of the current understanding of the molecular and genetic mechanisms 

underlying plasticity relevant to climate change. By bringing 

ecological, evolutionary, physiological and molecular perspectives together, we hope to provide clear directives for future 

research and stimulate cross-disciplinary dialogue on the relevance of phenotypic plasticity under climate change.

Link: http://www.ncbi.nlm.nih.gov/pubmed/20970368

97. Nitschke CR, Innes JL. 2008. A tree and climate 

assessment tool for modelling ecosystem response to climate 

change. Ecological Modelling 210:263-277.

Type: Journal 

Geographic Area: Western Canada

Keywords: climate change, regeneration, modelling, resilience, ecosystem, forest management, vulnerability, niche 

theory

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 35

from fire may govern post-fire recruitment response, we suggest that parental climate is the key selective force determining 

dormancy-breaking threshold temperatures, ensuring inter-fire 

seed bank persistence.

100. Opdam P, Wascher D. 2004. Climate change 

meets habitat fragmentation: linking landscape and biogeographical scale levels in research and conservation. Biological 

Conservation 117:285-297.

Type: Journal 

Geographic Area: Global

Keywords: climate change, habitat fragmentation, metapopulation dynamics, geographical range, habitat networks

Abstract: Climate change and habitat fragmentation are 

considered key pressures on biodiversity. In this paper we 

explore the potential synergetic effects between these factors. We argue that processes at two levels of spatial scale 

interact: the metapopulation level and the species range level. Current concepts of spatially dynamic metapopulations 

and species ranges are consistent, and integration improves 

our understanding of the interaction of landscape level and 

geographical range level processes. In landscape zones in 

which the degree of habitat fragmentation allows persistence, the shifting of ranges is inhibited, but not blocked. 

In areas where the spatial cohesion of the habitat is below 

the critical level of metapopulation persistence, the expansion of ranges will be blocked. An increased frequency of 

large-scale disturbances caused by extreme weather events 

will cause increasing gaps and an overall contraction of the 

distribution range, particularly in areas with relatively low 

levels of spatial cohesion. Taking into account the effects of 

climate change on metapopulations, habitat distribution and 

land use changes, future biodiversity research and conservation strategies are facing the challenge to re-orient their focus and scope by integrating spatially and conceptually more 

dynamic aspects at the landscape level.

Link: http://research.eeescience.utoledo.edu/lees/Teaching/

EEES4760_07/Opdam.PDF

101. Oregon Forest Resources Institute. 2006.

Forests, carbon and climate change: a synthesis of science findings. Portland (OR): Oregon Forest Resources Institute and 

Oregon State University. 192 p. 

Type: Book

Geographic Area: Western USA

Compilers’ Keywords: forest management, carbon dioxide, 

scale, strategies

Preface: The Oregon Forest Resources Institute (OFRI) commissioned this book, a synthesis of science findings on the 

relationships between forests, atmospheric carbon and climate 

change. While there is not scientific consensus about all the 

And, if we heed their warnings, caring for the birds can help us 

protect ourselves and the future of the world we share. Amid 

mounting concerns over accelerating global climate change, 

Audubon looked to the birds to determine if and how these 

sensitive creatures might be responding to changes here in the 

continental US. Birders have long reported surprising sightings of species far north of expected ranges. But are the reports 

significant? If so, are they connected to documented changes 

in our climate? Analyses of four decades of Christmas Bird 

Count data provide some answers. The results confirm what 

bird lovers have long suspected. Findings summarized in the 

pages that follow offer a look at forty years of change, a peek 

at what the future likely holds in one part of our nation, and 

an urgent message of warning from the birds—a message we 

would be wise to heed.

Link: http://web4.audubon.org/news/pressroom/bacc/pdfs/

birds%20and%20climate%20report.pdf

99. Ooi MKJ, Auld TD, Denham AJ. 2011. Projected 

soil temperature increase and seed dormancy response along an 

altitudinal gradient: implications for seed bank persistence under climate change. Plant and Soil 353:289-303.

Type: Journal 

Geographic Area: Australia

Keywords: Fabaceae, fire, heat wave, local adaptation, maternal effects, physiological dormancy, seed ecology, legume

Abstract: Understanding the mechanistic effects of climate 

change on species key life-history stages is essential for predicting ecological responses. In fire-prone regions, long-term seed 

banks allow post-fire recovery and persistence of plant populations. For physically dormant species, seed bank longevity 

depends on the maintenance of dormancy which is controlled 

primarily by temperature. Successful inter-fire recruitment is 

rare and dormancy loss between fires produces a net loss to the 

seed bank. We assessed whether temperature increases related 

to climate change can affect seed dormancy and, potentially, 

seed bank longevity. We quantified the relationship between 

air temperatures and soil temperatures. Seeds of two shrub species, from four populations along an altitudinal gradient, were 

then exposed to a range of soil temperatures calculated to occur at the end of the 21st century, using projected mean and 

heat wave scenarios. Alterations to dormancy were assessed 

via germination. For every 1 °C increase in air temperature, 

associated soil temperature increased by 1.5 °C. Mean temperature increase had no effect on seed dormancy. However, 

future heat wave conditions produced soil temperatures that 

significantly increased dormancy loss. This impact was greatest in seeds from cooler, high elevation populations. Projected 

heat wave events produce conditions that provide a mechanism 

for seed bank compromise. Dormancy-breaking temperatures 

for each population were positively related to parental environment temperatures, indicating local adaptation. Whilst heat 

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
36 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Link: http://www.discoverlife.org/pa/or/polistes/pr/2010nsf_

macro/references/Parmesan2007.pdf

103. Polley HW, Briske DD, Morgan JA, Wolter K, 

Bailey DW, Brown JR. 2013. Climate change and North 

American rangelands: trends, projections, and implications. 

Rangeland Ecology & Management 66:493-511.

Type: Journal 

Geographic Area: North America

Keywords: atmospheric CO2, atmospheric warming, climate 

variability, greenhouse gases, livestock production, precipitation patterns

Abstract: The amplified ‘‘greenhouse effect’’ associated 

with increasing concentrations of greenhouse gases has 

increased atmospheric temperature by 1 °C since industrialization (around 1750), and it is anticipated to cause an additional 2 °C increase by mid-century. Increased biospheric 

warming is also projected to modify the amount and distribution of annual precipitation and increase the occurrence of 

both drought and heat waves. The ecological consequences 

of climate change will vary substantially among ecoregions 

because of regional differences in antecedent environmental 

conditions; the rate and magnitude of change in the primary 

climate change drivers, including elevated carbon dioxide 

(CO2), warming and precipitation modification; and no additive effects among climate drivers. Elevated atmospheric 

CO2 will directly stimulate plant growth and reduce negative 

effects of drying in a warmer climate by increasing plant 

water use efficiency; however, the CO2 effect is mediated 

by environmental conditions, especially soil water availability. Warming and drying are anticipated to reduce soil water 

availability, net primary productivity, and other ecosystem 

processes in the southern Great Plains, the Southwest, and 

northern Mexico, but warmer and generally wetter conditions 

will likely enhance these processes in the northern Plains and 

southern Canada. The Northwest will warm considerably, 

but annual precipitation is projected to change little despite 

a large decrease in summer precipitation. Reduced winter 

snowpack and earlier snowmelt will affect hydrology and 

riparian systems in the Northwest. Specific consequences 

of climate change will be numerous and varied and include 

modifications to forage quantity and quality and livestock 

production systems, soil C content, fire regimes, livestock 

metabolism, and plant community composition and species 

distributions, including range contraction and expansion of 

invasive species. Recent trends and model projections indicate continued directional change and increasing variability 

in climate that will substantially affect the provision of ecosystem services on North American rangelands.

Link: http://www.climate-eval.org/sites/default/files/news/

study%20on%20effects%20of%20climate%20change%20

on%20north%20american%20rangelands.pdf

causes and implications of global climate change and the role 

of human activities, there is agreement that the relationships between forests and carbon, carbon and climate, and climate and 

forests are important and need to be better understood. It is also 

clear that Oregon is a forest-rich state, poised with opportunities 

for forests, forestry and forest product enterprises to contribute 

toward maintaining a livable climate.

Link: http://tongassmonitoring.com/pdf/Forests-CarbonClimateChange-2006.pdf#page=39

102. Parmesan C. 2007. Influences of species, latitudes 

and methodologies on estimates of phenological response to 

global warming. Global Change Biology 13:1860-1872.

Type: Journal 

Geographic Area: Northern Hemisphere

Keywords: animal, climate change, global warming, insect–

plant interactions, latitude, meta-analysis, phenology, plant, 

temperature, trophic interactions

Abstract: New analyses are presented addressing the global 

impacts of recent climate change on phenology of plant and 

animal species. A meta-analysis spanning 203 species was conducted on published datasets from the northern hemisphere. 

Phenological response was examined with respect to two factors: distribution of species across latitudes and taxonomic affiliation or functional grouping of target species. Amphibians 

had a significantly stronger shift toward earlier breeding than 

all other taxonomic/functional groups, advancing more than 

twice as fast as trees, birds and butterflies. In turn, butterfly 

emergence or migratory arrival showed three times stronger advancement than the first flowering of herbs, perhaps portending 

increasing asynchrony in insect–plant interactions. Response 

was significantly stronger at higher latitudes where warming 

has been stronger, but latitude explained < 4% of the variation. 

Despite expectation, latitude was not yet an important predictor 

of climate change impacts on phenology. The only two previously published estimates of the magnitude of global response 

are quite different: 2.3 and 5.1 days decade-1 advancement. The 

scientific community has assumed this difference to be real and 

has attempted to explain it in terms of biologically relevant 

phenomena: specifically, differences in distribution of data 

across latitudes, taxa or time periods. Here, these and other possibilities are explored. All analyses indicate that the difference 

in estimated response is primarily due to differences between 

the studies in criteria for incorporating data. It is a clear and 

automatic consequence of the exclusion by one study of data 

on “stable” (nonresponsive) species. Once this is accounted for, 

the two studies support each other, generating similar conclusions despite analyzing substantially no overlapping datasets. 

Analyses here on a new expanded dataset estimate an overall 

spring advancement across the northern hemisphere of 2.8 days 

decade-1. This is the first quantitative analysis showing that 

data-sampling methodologies significantly impact global (synthetic) estimates of magnitude of global warming response.

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 37

Compilers’ Keywords: climate change, endangered species, 

body size

Synopsis: This paper describes how climate variation in time 

and space can constrain community structure on a global scale. 

We explore body size scaling and the energetic consequences 

in terms of absorbed mass and energy and expended mass and 

energy. We explain how morphology, specific physiological 

properties, and temperature dependent behaviors are key variables that Link individual energetics to population dynamics 

and community structure. This paper describes an integrated 

basic principles model for mammal energetics and extends the 

model to bird energetics. The model additions include molar 

balance models for the lungs and gut. The gut model couples 

food ingested to respiratory gas exchanges and evaporative 

water loss from the respiratory system. We incorporate a novel 

thermoregulatory model that yields metabolic calculations as a 

function of temperature. The calculations mimic empirical data 

without regression. We explore the differences in the quality 

of insulation between hair and feathers with our porous media 

model for insulation. For mammals ranging in size from mice 

to elephants we show that calculated metabolic costs are in 

agreement with experimental data. We also demonstrate how 

we can do the same for birds ranging in size from hummingbirds to ostriches. We show the impact of changing posture 

and changing air temperatures on energetic costs for birds and 

mammals. We demonstrate how optimal body size that maximizes the potential for growth and reproduction changes with 

changing climatic conditions and with diet quality. Climate 

and diet may play important roles in constraining community 

structure (collection of functional types of different body sizes) 

at local and global scales. Thus, multiple functional types may 

coexist in a locality in part because of the temporal and spatial 

variation in climate and seasonal food variation. We illustrate 

how the models can be applied in a conservation and biodiversity context to a rare and endangered species of parrot, the 

Orange-bellied Parrot of Australia and Tasmania.

106. Pretzsch H, Biber P, Schutze G, Uhl E, Rotzer 

T. 2014. Forest stand growth dynamics in Central Europe have 

accelerated since 1870. Nature Communications 5:1-10.

Type: Journal 

Geographic Area: Central Europe

Compilers’ Keywords: climate change, tree growth, projections, forest management

Abstract: Forest ecosystems have been exposed to climate 

change for more than 100 years, whereas the consequences 

on forest growth remain elusive. Based on the oldest existing experimental forest plots in Central Europe, we show 

that, currently, the dominant tree species Norway spruce and 

European beech exhibit significantly faster tree growth (+32 to 

77%), stand volume growth (+10 to 30%) and standing stock 

accumulation (+6 to 7%) than in 1960. Stands still follow 

104. Ponce Campos GE, Moran MS, Huete A, 

Zhang Y, Bresloff C, Huxman TE, Eamus D, Bosch 

DD, Buda AR, Gunter SA, Scalley TH, Kitchen SG, 

McClaran MP, McNab WH, Montoya DS, Morgan 

JA, Peters DP, Sadler EJ, Seyfried MS, Starks 

PJ. 2013. Ecosystem resilience despite large-scale altered 

hydroclimatic conditions. Nature 494:349-352.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: climate change, drought, 20th century, 21st century, plants, Poaceae, rain, trees, water

Abstract: Climate change is predicted to increase both 

drought frequency and duration, and when coupled with substantial warming, will establish a new hydroclimatological 

model for many regions. Large-scale, warm droughts have recently occurred in North America, Africa, Europe, Amazonia 

and Australia, resulting in major effects on terrestrial ecosystems, carbon balance and food security. Here we compare the 

functional response of above-ground net primary production 

to contrasting hydroclimatic periods in the late twentieth century (1975–1998), and drier, warmer conditions in the early 

twenty-first century (2000–2009) in the Northern and Southern 

Hemispheres. We find a common ecosystem water-use efficiency (WUE(e): above-ground net primary production/evapotranspiration) across biomes ranging from grassland to forest 

that indicates an intrinsic system sensitivity to water availability across rainfall regimes, regardless of hydroclimatic conditions. We found higher WUE(e) in drier years that increased 

significantly with drought to a maximum WUE(e) across all 

biomes; and a minimum native state in wetter years that was 

common across hydroclimatic periods. This indicates biomescale resilience to the interannual variability associated with 

the early twenty-first century drought—that is, the capacity to 

tolerate low, annual precipitation and to respond to subsequent 

periods of favourable water balance. These findings provide 

a conceptual model of ecosystem properties at the decadal 

scale applicable to the widespread altered hydroclimatic conditions that are predicted for later this century. Understanding 

the hydroclimatic threshold that will break down ecosystem 

resilience and alter maximum WUE(e) may allow us to predict 

land-surface consequences as large regions become more arid, 

starting with water-limited, low-productivity grasslands.

Link: http://www.ncbi.nlm.nih.gov/pubmed/23334410 

105. Porter WP, Budaraju S, Stewart WE, 

Ramankutty N. 2000. Calculating climate effects on birds 

and mammals: impacts on biodiversity, conservation, population parameters, and global community structure. American 

Zoologist 10:597-630.

Type: Journal 

Geographic Area: Global

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
38 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

108. Rehfeldt GE, Ferguson DE, Crookston NL. 

2009. Aspen, climate, and sudden decline in western USA.

Forest Ecology and Management 258:2353-2364.

Type: Journal 

Geographic Area: Western USA

Keywords: bioclimate models, sudden aspen decline, global 

warming impacts, climate responses, climatic niche

Abstract: A bioclimate model predicting the presence or 

absence of aspen, Populus tremuloides, in western USA from 

climate variables was developed by using the Random Forests 

classification tree on Forest Inventory data from about 118, 

000 permanent sample plots. A reasonably parsimonious 

model used eight predictors to describe aspen’s climate profile. Classification errors averaged 4.5%, most of which were 

errors of commission. The model was driven primarily by 

three variables: an annual dryness index, the ratio of summer 

to annual precipitation, and an interaction of growing season 

precipitation with the summer–winter temperature differential. 

Projecting the contemporary climate profile into the future climate provided by three General Circulation Models and two 

scenarios (SRES A2 and either B1 or B2) suggested that the 

area occupied by the profile should diminish rapidly over the 

course of the century, 6–41% by the decade surrounding 2030, 

40–75% for that surrounding 2060, and 46–94% for 2090. 

The relevance of the climate profile to understanding climatebased responses is illustrated by relating trends in climate to 

the recent incidence of sudden aspen dieback that has plagued 

portions of the aspen distribution. Of the eight variables in the 

profile, four reached extreme values during 2000–2003, the period immediately preceding the appearance of damage in aerial 

surveys.

Link: http://www.treesearch.fs.fed.us/pubs/33823

109. Rietkerk M, Dekker SC, de Ruiter PC, van de 

Koppel J. 2004. Self-organized patchiness and catastrophic 

shifts in ecosystems. Science 305:1926-1929.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: climate change, range shifts, ecosystem management

Abstract: Unexpected sudden catastrophic shifts may occur 

in ecosystems, with concomitant losses or gains of ecological 

and economic resources. Such shifts have been theoretically 

attributed to positive feedback and biostability of ecosystem 

states. However, verifications and predictive power with respect to catastrophic responses to a changing environment 

are lacking for spatially extensive ecosystems. This situation 

impedes management and recovery strategies for such ecosystems. Here, we review recent studies on various ecosystems 

that link self-organized patchiness to catastrophic shifts between ecosystem states.

similar general allometric rules, but proceed more rapidly 

through usual trajectories. As forest stands develop faster, tree 

numbers are currently 17-20% lower than in past same-aged 

stands. Self-thinning lines remain constant, while growth rates 

increase indicating the stock of resources have not changed, 

while growth velocity and turnover have altered. Statistical 

analyses of the experimental plots, and application of an 

ecophysiological model, suggest that mainly the rise in temperature and extended growing seasons contribute to increased 

growth acceleration, particularly on fertile sites.

Link: http://www.ncbi.nlm.nih.gov/pubmed/25216297

107. Pyke CR, Fischer DT. 2005. Selection of bioclimatically representative biological reserve systems under climate 

change. Biological Conservation 121:429-441.

Type: Journal 

Geographic Area: California, USA

Keywords: systematic conservation planning, climate 

change, habitat loss, bioclimatic representation, integer linearprogramming, vernal pools

Abstract: Biological reserves are intended to protect species, communities, and ecosystems in human-dominated 

landscapes. However, existing protected areas represent only 

relatively small, geographically biased samples of species and 

habitats. Climate change and habitat loss can exacerbate these 

biases and the net result is a small, skewed subset of historic 

environmental conditions. We developed a general model to 

improve the representation of environmental conditions across 

the range of at-risk species or any other elements targeted for 

conservation. We implemented the model as an integer linearprogramming problem to select additional areas to complement 

existing reserves and create new portfolios that are bioclimatically representative across a range of climatic scenarios. We 

demonstrated the use of the model for a small dataset including two hydrologic variables across the range of five species 

of fairy shrimp (Anostraca) in the Central Valley ecoregion of 

California, USA under three climate scenarios. The bioclimatic 

representation model identified solutions that meet biodiversity representation goals and substantially improve bioclimatic 

representation at minimal additional cost in terms of total land 

selected for a conservation portfolio. Additional constraints 

rewarding bioclimatic representation under two conflicting climate scenarios resulted in only a small decrease in the 

performance of solutions with respect to current climate. We 

conclude that this model provides a general tool for improving 

bioclimatic representation, and results from the Central Valley 

case study suggest an encouraging, testable hypothesis that climatically robust bioclimatic representation can be achieved at 

negligible marginal costs.

Link: http://www.geog.ucsb.edu/~fischer/PykeFischer2005.

pdf

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 39

common garden in Petawawa, Ontario, Canada. Forest Ecology 

and Management 242:636-647.

Type: Journal 

Geographic Area: Ontario, Canada

Keywords: adaptation, climate change, GCM, Pinus banksiana, provenance, radial growth, transfer functions

Abstract: The effects of seed transfer and climate change 

on the width and basal area of tree rings were studied in 21 

provenances of jack pine (Pinus banksiana Lamb.) grown in 

a common-garden plantation in Petawawa, Ontario, Canada. 

Seed-source origin significantly influences both mean treering width and mean annual basal area increment over a 

25-year growth period (1975-1999). Temperature and precipitation transfer functions were developed to predict width 

and basal area of tree rings of the jack pine populations. The 

best predictors of growth were the transfer distances of mean 

annual maximum daily temperature and annual precipitation between the plantation site and the seed origins. Radial 

growth of the jack pine populations was mainly related to 

temperature at seed origin and, to a lesser degree, to precipitation at seed origin. Extension of the transfer functions to three 

sets of independent data revealed signification correlations 

between estimated and predicted mean radial growth characteristics. Seed sources of jack pine originating from warmer 

and drier climates than that of the plantation site in Petawawa 

had slightly higher mean ring widths and basal areas than 

the local populations. The application of different climate 

change scenarios derived from general circulation models to 

the developed transfer functions indicated that radial growth 

of jack pine may decline only if significant climate changes 

occur, which might not happen before the mid-21st century. 

Both a higher radial growth of southern seed sources and potential negative effect of a significant temperature increase 

and precipitation decrease in future suggest restricting the 

northward transfer of southern seed sources to less than 1 

degree latitude. However, provenance specific differences in 

survivorship, frost- and disease-resistance, and cone serotiny 

should also be taken into consideration. 

112. Schoettle AW, Sniezko R, Burns KS. 2009.

Sustaining Pinus flexilis ecosystems of the southern Rocky 

Mountains (USA) in the presence of Cronartium ribicola and 

Dendroctonus ponderosae in a changing climate. In: Noshad D, 

Noh Eun W, King J, Sniezko R, editors. Breeding and Genetic 

Resources of Five-Needle Pines. Seoul, Republic of South 

Korea: Korea Forest Research Institute. p 63-65.

Type: Conference Paper

Geographic Area: Northwestern USA

Compilers’ Keywords: limber pine, genetic variation, forest management, forest health, pest management, forest 

regeneration

Link: http://www.sciencemag.org/content/305/5692/1926.

short

110. Robbirt KM, Davy AJ, Hutchings MJ, Roberts 

DL. 2011. Validation of biological collections as a source 

of phenological data for use in climate change studies: a case 

study with the orchid Ophrys sphegodes. Journal of Ecology 

99:235-241.

Type: Journal 

Geographic Area: Southern and Central Europe

Keywords: biological collections, climate change, flowering 

time, herbarium specimens, natural history collections, Ophrys 

sphegodes, Orchidaceae, phenology, spring, temperature

Summary: The scarcity of reliable long-term phenological 

data has severely hindered the study of the responses of species 

to climate change. Biological collections in herbaria and museums are potential sources of long term data for such study, but 

their use for this purpose needs independent validation. Here 

we report a rigorous test of the validity of using herbarium 

specimens for phenological studies, by comparing relationships between climate and time of peak flowering derived from 

herbarium records and from direct field-based observations, 

for the terrestrial orchid Ophrys sphegodes. We examined herbarium specimens of O. sphegodes collected between 1848 and 

1958, and recorded peak flowering time directly in one population of O. sphegodes between 1975 and 2006. The response 

of flowering time to variation in mean spring temperature 

(March–May) was virtually identical in both sets of data, even 

though they covered different periods of time which differ in 

extent of anthropogenic temperature change. In both cases 

flowering was advanced by about 6 days per °C rise in average 

spring temperature. The proportion of variation in flowering 

time explained by spring temperature was lower in the herbarium record than in direct field observations. It is likely that 

some of the additional variation was due to geographical variation in collection site, as flowering was significantly earlier at 

more westerly sites, which have had warmer springs, over their 

range of 3.44° longitude. Predictions of peak flowering time 

based on the herbarium data corresponded closely with observed peak flowering times in the field, indicating that flowering response to temperature had not altered between the two 

separate periods over which the herbarium and field data were 

collected. These results provide the first direct validation of the 

use of herbarium collections to examine the relationships between phenology and climate when field-based observational 

data are not available.

Link: http://www.drdavidroberts.com/docs/Robbirt_

etal_2011_Ophrys_Climate_Change.pdf

111. Savva Y, Denneler B, Koubaa A, Tremblay F, 

Bergeron Y, Tjoelker MG. 2007. Seed transfer and climate change effects on radial growth of jack pine populations in a 

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
40 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

suggest that there is a strong need to intensify monitoring 

efforts and to develop additional conservation measures for 

populations in the most vulnerable units. Also, our results 

call for continued transnational actions for genetic conservation of European forest trees, including the establishment of 

dynamic conservation populations outside the current species distribution ranges within European assisted migration 

schemes.

Link: http://www.ncbi.nlm.nih.gov/pubmed/24273066

114. Schwartz MW, Iverson LR, Prasad AM, 

Matthews SN, O’Connor RJ. 2006. Predicting extinctions as a result of climate change. Ecology 87:1611-1615.

Type: Journal 

Geographic Area: North America

Keywords: climate and environmental models, climate 

change, distribution breadth, eastern United States, endemic, 

extinction, prediction uncertainty, regression tree, vulnerability

Abstract: Widespread extinction is a predicted ecological 

consequence of global warming. Extinction risk under climate 

change scenarios is a function of distribution breadth. Focusing 

on trees and birds of the eastern United States, we used joint 

climate and environment models to examine fit and climate 

change vulnerability as a function of distribution breadth. We 

found that extinction vulnerability increases with decreasing 

distribution size. We also found that model fit decreases with 

decreasing distribution size, resulting in high prediction uncertainty among narrowly distributed species. High prediction 

uncertainty creates a conservation dilemma in that excluding 

these species under predicts extinction risk and favors mistaken 

inaction on global warming. By contrast, including narrow endemics results in over-predicting extinction risk and promotes 

mistaken inaction on behalf of individual species prematurely 

considered doomed to extinction.

Link: http://kb.osu.edu/dspace/bitstream/handle/1811/49027/

fac_IversonL_Ecology_2006_87_7.pdf?sequence=1

115. Seager R, Ting M, Held I, Kushnir Y, Lu J, 

Vecchi G, Huang HP, Harnik N, Leetmaa A, Lau 

NC, Li C, Velez J, Naik N. 2007. Model projections of 

an imminent transition to a more arid climate in southwestern 

North America. Science 316:1181-1184.

Type: Journal 

Geographic Area: Southwestern USA

Compilers’ Keywords: hydrology, climate change

Abstract: How anthropogenic climate change will affect hydroclimate in the arid regions of southwestern North America 

has implications for the allocation of water resources and the 

course of regional development. Here we show that there is a 

broad consensus among climate models that this region will 

dry in the 21st century and that the transition to a more arid 

Abstract: Limber pine, Pinus flexilis James, is characterized by a patchy distribution that displays metapopulation 

dynamics and spans a broad latitudinal and elevational range 

in North America. In the southern Rocky Mountains, limber 

pine grows from below the forest-grassland ecotone up to 

the forest-alpine ecotone, from ~1600 m above sea level in 

the short grass steppe to > 3300 m at the continental divide. 

In this region, limber pine’s altitudinal range is wider than 

any of its co-occurring tree species. Limber pine ecosystems 

serve a variety of important ecological roles, such as (1) occupying and stabilizing dry habitats, (2) defining ecosystem 

boundaries (treelines), (3) being among the first tree species 

to colonize a site after fire, (4) facilitating the establishment 

of late successional species and (5) providing diet and habitat 

for animals.

Link: http://www.treesearch.fs.fed.us/pubs/34716

113. Schueler S, Falk W, Koskela J, Lefevre F, 

Bozzano M, Hubert J, Kraigher H, Longauer R, 

Olrik DC. 2014. Vulnerability of dynamic genetic conservation units of forest trees in Europe to climate change. Global 

Change Biology 20:1498-1511.

Type: Journal 

Geographic Area: Europe

Keywords: climate niche modelling, favourability function, 

forest genetic resources, gap analysis, genetic conservation 

unit, velocity of climate change

Abstract: A transnational network of genetic conservation 

units for forest trees was recently documented in Europe 

aiming at the conservation of evolutionary processes and the 

adaptive potential of natural or man-made tree populations. 

In this study, we quantified the vulnerability of individual 

conservation units and the whole network to climate change 

using climate favourability models and the estimated velocity of climate change. Compared to the overall climate 

niche of the analysed target species populations at the warm 

and dry end of the species niche are underrepresented in 

the network. However, by 2100, target species in 33-65% 

of conservation units, mostly located in southern Europe, 

will be at the limit or outside the species’ current climatic 

niche as demonstrated by favourabilities below required 

model sensitivities of 95%. The highest average decrease in 

favourabilities throughout the network can be expected for 

coniferous trees although they are mainly occurring within 

units in mountainous landscapes for which we estimated 

lower velocities of change. Generally, the species-specific 

estimates of favourabilities showed only low correlations to 

the velocity of climate change in individual units, indicating 

that both vulnerability measures should be considered for 

climate risk analysis. The variation in favourabilities among 

target species within the same conservation units is expected 

to increase with climate change and will likely require a prioritization among co-occurring species. The present results 

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 41

will not be adapted to future climates. We estimated the risk 

of maladaptation of current populations of coastal Douglas-fir 

(Pseudotsuga menziesii var. menziesii) to future climates as the 

proportion of no overlap between two normal distributions 

where the means and genetic variances of current and future 

populations are determined from genecological models derived 

from seedling common garden studies. The risk of maladaptation was large for most traits when compared with the risk associated with current transfers within seed zones, particularly 

for the more drastic climate change scenario. For example, the 

proportion of no overlap for a composite trait representing bud 

set, emergence, growth, and root : shoot ratio was as high as 

0.90. We recommend augmenting within-population variation 

by mixing local populations with some proportion of populations from lower elevations and further south. Populations 

expected to be adapted to climates a century from now come 

from locations as far down in elevation as 450–1130 m and as 

far south in latitude as 1.8–4.9°.

118. Strzepek K, Yohe G, Neumann J, Boehlert B. 

2010. Characterizing changes in drought risk for the United 

States from climate change. Environmental Research Letters 

5:044012.

Type: Journal 

Geographic Area: USA

Keywords: drought, climate change, drought severity indices, 

uncertainty

Abstract: The effect of climate change on the frequency 

and intensity of droughts across the contiguous United States 

over the next century is assessed by applying Standardized 

Precipitation Indices and the Palmer Drought Severity Index 

to the full suite of 22 Intergovernmental Panel on Climate 

Change General Circulation Models for three IPCC-SRES 

emissions scenarios (B1, A1B, and A2 from the Special Report 

on Emissions Scenarios (SRES) listed in order of their emissions through 2100 from high to low). The frequency of meteorological drought based on precipitation alone is projected 

to increase in some parts of the U.S., for example the southwestern states, and decrease in others. Hydrological drought 

frequencies based on precipitation and temperature are projected to increase across most of the country, however, with 

very substantial and almost universally experienced increases 

in drought risk by 2050. For both measures, the southwestern 

U.S. and the Rocky Mountain states are projected to experience the largest increases in drought frequency, but these areas may be able to exploit existing excess storage capacity. 

Drought frequencies and uncertainties in their projection tend 

to increase considerably over time and show a strong worsening trend along higher greenhouse gas emissions scenarios, 

suggesting substantial benefits for greenhouse gas emissions 

reductions.

Link: http://globalchange.mit.edu/files/document/

MITJPSPGC_Reprint_2010-14.pdf

climate should already be under way. If these models are correct, the levels of aridity of the recent multiyear drought or 

the Dust Bowl and the 1950s droughts will become the new 

climatology of the American Southwest within a time frame of 

years to decades.

Link: http://www.ncbi.nlm.nih.gov/pubmed/17412920

116. Soolanayakanahally RY, Guy RD, Silim SN, 

Song M. 2012. Timing of photoperiodic competency causes 

phenological mismatch in balsam poplar (Populus balsamifera

L.). Plant Cell and Environment 36:116-127.

Type: Journal 

Geographic Area: Canada

Keywords: bud flush, bud set, climate change, growing 

season length, height growth cessation, latitude, phenology, 

photoperiod

Abstract: Plant phenology is expected to be sensitive to climate warming. In boreal trees, spring flush is primarily temperature driven, whereas height growth cessation and autumn 

leaf senescence are predominantly controlled by photoperiod. 

Cuttings of 525 genotypes from the full range of balsam poplar 

were planted into two common gardens (Vancouver and Indian 

Head, Canada) at similar latitudes, but with differing winter 

temperatures and growing seasons. There was clinal variation 

in spring and, particularly, summer and fall phenology. Bud 

flush and, despite milder climate, bud set and leaf drop were 

earlier at Vancouver than at Indian Head by 44, 28 and 7 d, 

respectively. Although newly flushed growth is insensitive to 

photoperiod, many genotypes at both sites became competent 

before the summer solstice. At Vancouver, high-latitude genotypes set dormant terminal buds in mid-spring. Most other genotypes grew until midsummer or set bud temporarily and then 

experienced a second flush. In both gardens and in a growth 

chamber experiment, earlier bud set was associated with reduced height growth and higher root/shoot ratios. Shoots attained competency approximately 5 weeks after flushing, 

which would normally prevent dormancy induction before the 

solstice, but may be insufficient if spring advances by more 

than a few weeks.

Link: http://www.ncbi.nlm.nih.gov/pubmed/22702736

117. St. Clair JB, Howe GT. 2007. Genetic maladaptation of coastal Douglas-fir seedlings to future climates. Global 

Change Biology 13:1441-1454.

Type: Journal 

Geographic Area: Pacific Northwest USA

Keywords: climate change, forest trees, genecology, local adaptation, natural selection, precipitation, Pseudotsuga menziesii, 

quantitative traits, relative risk of maladaptation, temperature

Abstract: Climates are expected to warm considerably over 

the next century, resulting in expectations that plant populations 

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
42 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

across a substantial elevation range (1800–2700 m) and at both 

aggregate watershed and 90 m patch scales. Model results suggest that the variation in AET due to recharge timing is greatest for patches and years with moderate levels of precipitation 

or patches that receive substantial lateral moisture inputs. For 

a 3 °C warming scenario, the annual AET increases in some 

years due to warmer temperatures but decreases by as much 

as 40% in other years due to an earlier timing of snowmelt. 

These results help to clarify the conditions under which water 

availability for forests decreases and highlight scenarios that 

may lead to increased drought stress under a warming climate 

in snow-dominated mountain regions.

121. Tepe TL, Meretsky VJ. 2011. Forward-looking forest restoration under climate change—are U.S. nurseries ready? 

Restoration Ecology 19:295-298.

Type: Journal 

Geographic Area: USA

Keywords: forest restoration, planning, restoration planting, 

state nurseries

Abstract: The pace of climate change suggests that restoration 

efforts once focused on past conditions should become more 

forward-looking. Suggestions for such restoration emphasize 

the use of a suite of species adapted to a range of possible future climates. In forest restoration, opportunities for forwardlooking restoration may be limited by the availability of suitable stock from state and commercial nurseries. Presently, most 

state nurseries have stock potentially suited to warmer climates 

than currently exist in their states. However, these nurseries are 

generally not actively incorporating information about climate 

change into their stocking choices and some see clear obstacles 

to providing such stock, particularly uncertainty about the future climate, and the existence of seed zones and other policies 

designed to protect locally adapted species genetics. As restoration ecologists adapt their methods to incorporate climate 

change, state nurseries should be involved in those discussions 

and may be important partners in outreach.

122. Thomas CD, Hill JK, Anderson BJ, Bailey S, 

Beale CM, Bradbury RB, Bulman CR, Crick HQP, 

Eigenbrod F, Griffiths HM, Kunin WE, Oliver TH, 

Walmsley CA, Watts K, Worsfold NT, Yardley T. 

2011. A framework for assessing threats and benefits to species responding to climate change. Methods in Ecology and 

Evolution 2:125-142.

Type: Journal 

Geographic Area: Britain, Global

Keywords: biodiversity, climate envelope, distribution, global 

warming, IUCN, policy, risk assessment, species conservation

Summary: Current national and international frameworks 

for assessing threats to species have not been developed in the 

119. Swift K, Ran S. 2012. Successional responses to 

natural disturbance, forest management, and climate change 

in British Columbia’s Forests. Journal of Ecosystems and 

Management 13:1-23.

Type: Journal 

Geographic Area: Western Canada

Keywords: successional pathways, climate change, natural 

disturbance, fire, insects, diseases

Abstract: Natural and human-induced disturbance such as 

wildfire, insect and disease outbreak, windthrow, and forest harvesting are important drivers for forest renewal, postdisturbance stand structure, and ecosystem function. Each 

disturbance or combination of disturbances sets up a forest 

to proceed down a certain successional pathway in terms of 

structure and function. Using the context of Ecoprovinces and 

Ecosystem Types, successional pathways of a variety of ecosystems found in British Columbia are briefly described, and 

the ways in which forest management practices have affected 

those pathways are discussed. This Extension Note also describes how projected changes in temperature and precipitation 

may also affect these natural disturbance drivers. The information contained in this article is based on a larger synthesis 

report that is available in FORREX Series 28 and is designed 

to facilitate further conservation around building resistant and 

resilient forests for the future. 

Link: http://www.jem.forrex.org/forrex/images/PDFs/JEM_

VOL_13_NO_1.pdf#page=62

120. Tague C, Peng H. 2013. The sensitivity of forest water use to the timing of precipitation and snowmelt recharge in 

the California Sierra: implications for a warming climate. Journal 

of Geophysical Research: Biogeosciences 118:875-887.

Type: Journal 

Geographic Area: California, USA

Compilers’ Keywords: climate change, hydrologic change

Abstract: Forest responses to warming, in the absence of 

changes in vegetation structure, reflect the balance between 

the increased atmospheric demand for water and changes in 

water availability. This study uses a coupled hydroecologic 

model applied to a snow-dominated mountain watershed to 

demonstrate how complex topography and interannual variation in climate drivers combine to alter the balance between 

moisture availability and energy demand. We focus specifically on how often and under what conditions changes in the 

timing of soil water recharge as precipitation or snowmelt are a 

significant control on forest actual evapotranspiration (AET) in 

the Central California Sierra. We show that while interannual 

variation in precipitation is the dominant control on interannual variation in AET, how much of that recharge accumulates 

as a seasonal snowpack can act as a second-order control. 

This sensitivity of AET to snow accumulation and melt occurs 

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 43

to increased water vapor in the atmosphere. Hence, storms, 

whether individual thunderstorms, extratropical rain or snow 

storms, or tropical cyclones, supplied with increased moisture, produce more intense precipitation events. Such events 

are observed to be widely occurring, even where total precipitation is decreasing: “it never rains but it pours!” This increases the risk of flooding. The atmospheric and surface energy 

budget plays a critical role in the hydrological cycle, and also 

in the slower rate of change that occurs in total precipitation 

than total column water vapor. With modest changes in winds, 

patterns of precipitation do not change much, but result in dry 

areas becoming drier (generally throughout the subtropics) 

and wet areas becoming wetter, especially in the mid- to high 

latitudes: the “rich get richer and the poor get poorer.” This 

pattern is simulated by climate models and is projected to continue into the future. Because, with warming, more precipitation occurs as rain instead of snow and snow melts earlier, 

there is increased runoff and risk of flooding in early spring, 

but increased risk of drought in summer, especially over continental areas. However, with more precipitation per unit of 

upward motion in the atmosphere, i.e., “more bang for the 

buck,” atmospheric circulation weakens, causing monsoons 

to falter. In the tropics and subtropics, precipitation patterns 

are dominated by shifts as sea surface temperatures change, 

with El Niño a good example. The volcanic eruption of Mount 

Pinatubo in 1991 led to an unprecedented drop in land precipitation and runoff, and to widespread drought, as precipitation shifted from land to oceans and evaporation faltered, 

providing lessons for possible geoengineering. Most models 

simulate precipitation that occurs prematurely and too often, 

and with insufficient intensity, resulting in recycling that is too 

large and a lifetime of moisture in the atmosphere that is too 

short, which affects runoff and soil moisture.

Link: http://envsci.rutgers.edu/~toine379/extremeprecip/

papers/trenberth_2011.pdf

124. Vandersteen W. 2011. Detecting gene expression 

profiles associated with environmental stressors within an ecological context. Molecular Ecology 20:1322-1323.

Type: Journal 

Geographic Area: Global

Keywords: adaptation, climate change, conservation biology, 

genomics ⁄ proteomics transcriptomics

Abstract: Prior to the development of any conservation strategies to mitigate deleterious impacts of environmental change 

and contamination, there must be a method to make meaningful predictions of the effect of environmental change on 

organisms. Assessment of relative transcriptomic expression 

patterns can provide a link between the environment and the 

physiological response of the organism by identifying genes 

that respond to environmental stressors; this information could 

also assist in teasing apart the molecular basis of toxicological 

context of climate change, and are not framed in a way that recognizes new opportunities that arise from climate change. The 

framework presented here separates the threats and benefits of 

climate change for individual species. Threat is assessed by 

the level of climate-related decline within a species’ recently 

occupied (e.g., pre-1970s) historical distribution, based on 

observed (e.g., repeat census) and⁄or projected changes (e.g., 

modelled bioclimate space). Benefits are assessed in terms of 

observed and⁄ or projected increases outside the recently occupied historical range. Exacerbating factors (e.g., small population size, low dispersal capacity) that might increase levels 

of threat or limit expansion in response to climate change 

are taken into consideration within the framework. Protocols 

are also used to identify levels of confidence (and hence research and⁄or monitoring needs) in each species’ assessment. 

Observed and projected changes are combined into single 

measures of expected decline and increase, together with associated measures of confidence. We weight risk classifications towards information that is most certain. Each species is 

then placed in one of six categories (high risk, medium risk, 

limited impact, equivalent risks & benefits, medium benefit, 

high benefit) reflecting whether climate change is expected 

(or has been observed) to cause net declines or increases in 

the region considered, based on the balance of benefits and 

threats. We illustrate the feasibility of using the framework by 

applying it to (i) all British butterflies (N = 58 species) and (ii) 

an additional sample of British species: 18 species of plants, 

bats, birds and beetles. Our framework assesses net declines 

and increases associated with climate change, for individual 

species. It could be applied at any scale (regional, continental 

or global distributions of species), and complements existing 

conservation assessment protocols such as red-listing. Using 

observed and projected population and/or range data, it is feasible to carry out systematic conservation status assessments 

that inform the development of monitoring, adaptation measures and conservation management planning for species that 

are responding to climate change.

Link: http://www.researchgate.net/publication/229977974_A_

framework_for_assessing_threats_and_benefits_to_species_

responding_to_climate_change/file/79e4150c83bfe76a17.pdf

123. Trenberth KE. 2011. Changes in precipitation with 

climate change. Climate Research 47:123-138.

Type: Journal 

Geographic Area: Global

Keywords: climate change, precipitation, storms, drought, 

extremes, floods, geoengineering, climate models

Abstract: There is a direct influence of global warming on 

precipitation. Increased heating leads to greater evaporation 

and thus surface drying, thereby increasing the intensity and 

duration of drought. However, the water holding capacity of 

air increases by about 7% per 1 °C warming, which leads 

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
44 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

widespread species. Our global analysis of future climatic 

range change of common and widespread species shows that 

without mitigation, 57 ± 6% of plants and 34 ± 7 of animals 

are likely to lose ≥ 50% of their present climatic range by the 

2080s. With mitigation, however, losses are reduced by 60% 

if emissions peak in 2016 or 40% if emissions peak in 2030. 

Thus, our analyses indicate that without mitigation, large 

range contractions can be expected even amongst common 

and widespread species, amounting to a substantial global 

reduction in biodiversity and ecosystem services by the end 

of this century. Prompt and stringent mitigation, on the other 

hand, could substantially reduce range losses and buy up to 

four decades for climate change adaptation.

127. Warwell MV, Rehfeldt GE, Crookston NL. 

2010. Modeling species’ realized climatic niche space and 

predicting their response to global warming for several western 

forest species with small geographic distributions. In: Pye JM, 

Rauscher HM, Sands Y, Lee DC, Beatty JS, editors. Advances 

in threat assessment and their application to forest and rangeland management. Portland (OR): USDA Forest Service, 

Pacific Northwest and Southern Research Stations. General 

Technical Report PNW-GTR-802. p 171-182.

Type: Government Document

Geographic Area: Western North America

Keywords: bioclimatic models, climatic distributions, climatic niche, global warming, Random Forests, multiple-regression 

tree, response to climate change, narrow endemic

Abstract: The Random Forests multiple regression tree was 

used to develop an empirically based bioclimatic model of 

the presence absence of species occupying small geographic 

distributions in western North America. The species assessed 

were subalpine larch (Larix lyallii), smooth Arizona cypress 

(Cupressus arizonica ssp. glabra), Paiute cypress (syn. Piute cypress) (Cupressus arizonica ssp. nevadensis), and Macfarlane’s 

four-o’clock (Mirabilis macfarlanei). Independent variables 

included 33 simple expressions of temperature and precipitation and their interactions. These climate variables were 

derived from a spline climate model for the western United 

States that provides point estimates (latitude, longitude, and 

altitude). Analyses used presence-absence data largely from 

the Forest Inventory and Analysis, USDA Forest Service database. Overall errors of classification ranged from 1.39 percent for Macfarlane’s four-o’clock to 3.55 percent for smooth 

Arizona cypress. The mapped predictions of species occurrence using the estimated realized climatic niche space were 

more accurate than published range maps. The Hadley and 

Canadian general circulation models (scenario IS92a for 1 

percent increase GGa/year) were then used to illustrate the potential response of the species’ contemporary realized climatic 

niche space to climate change. Predictions were mapped at a 

1-km2 resolution. Concurrence between species’ geographic 

effects vs. physiological adaptation. Molecular responses to 

environmental stressors are probably not restricted to single 

or few genes, and therefore a more integrative approach is 

required to examine broad-scale patterns of transcriptomic response. To address this objective, Chapman et al. (2011) used 

machine learning tools to link the mechanisms of physiological response to environmental stress; although widely used 

in clinical applications, such as finding the genetic basis of 

diseases, ecological genomics applications of artificial neural networks are just beginning to emerge. Analyses such as 

these are important to help identify limitations on the adaptive capacity of organisms and to predict impacts of climate 

change, ocean acidification and anthropogenic contaminants 

on aquatic organisms.

125. Walther GR, Post E, Convey P, Menzel A, 

Parmesan C, Beebee TJC, Fromentin JM, HoeghGuldberg O, Bairlein F. 2002. Ecological responses to 

recent climate change. Nature 416:389-395.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: review, phenology, range shift

Abstract: There is now ample evidence of the ecological impacts of recent climate change, from polar terrestrial to tropical 

marine environments. The responses of both flora and fauna 

span an array of ecosystems and organizational hierarchies, 

from the species to the community levels. Despite continued 

uncertainty as to community and ecosystem trajectories under 

global change, our review exposes a coherent pattern of ecological change across systems. Although we are only at an early stage in the projected trends of global warming, ecological 

responses to recent climate change are already clearly visible.

Link: http://eebweb.arizona.edu/courses/Ecol206/

Walther%20et%20al%20Nature%202002.pdf

126. Warren R, VanDerWal JJ, Price J, Welbergen 

JA, Atkinson I, Ramirez-Villegas J, Osborn TJ, Jarvis 

A, Shoo LP, Williams SE, Lowe J. 2013. Quantifying 

the benefit of early climate change mitigation in avoiding biodiversity loss. Nature Climate Change May:1-5.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: species distribution, habitat contractions, emission reductions, plants, animals

Abstract: Climate change is expected to have significant 

influences on terrestrial biodiversity at all system levels, including species-level reductions in range size and abundance, 

especially amongst endemic species. However, little is known 

about how mitigation of greenhouse gas emissions could reduce biodiversity impacts, particularly amongst common and 

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 45

129. Westerling AL, Turner MG, Smithwick EAH, 

Romme WH, Ryan MG. 2011. Continued warming could 

transform Greater Yellowstone fire regimes by mid-21st century.

Proceedings of the National Academy of Science, Ecology, 

Environmental Sciences 108:13165-13170.

Type: Journal 

Geographic Area: Montana, Wyoming, USA

Compilers’ Keywords: climate change, wildfire regime

Abstract: Climate change is likely to alter wildfire regimes, 

but the magnitude and timing of potential climate-driven 

changes in regional fire regimes are not well understood. We 

considered how the occurrence, size, and spatial location of 

large fires might respond to climate projections in the Greater 

Yellowstone ecosystem (GYE) (Wyoming), a large wildland 

ecosystem dominated by conifer forests and characterized by 

infrequent, high-severity fire. We developed a suite of statistical models that related monthly climate data (1972–1999) to 

the occurrence and size of fires > 200 ha in the northern Rocky 

Mountains; these models were cross-validated and then used 

with downscaled (~12 km × 12 km) climate projections from 

three global climate models to predict fire occurrence and area 

burned in the GYE through 2099. All models predicted substantial increases in fire by midcentury, with fire rotation (the 

time to burn an area equal to the landscape area) reduced to 

< 30 y from the historical 100–300 y for most of the GYE. 

Years without large fires were common historically but are 

expected to become rare as annual area burned and the frequency of regionally synchronous fires increase. Our findings 

suggest a shift to novel fire–climate–vegetation relationships 

in Greater Yellowstone by midcentury because fire frequency 

and extent would be inconsistent with persistence of the current suite of conifer species. The predicted new fire regime 

would transform the flora, fauna, and ecosystem processes 

in this landscape and may indicate similar changes for other 

subalpine forests.

Link: http://www.greateryellowstone.org/uploads/GYE_and_

Fire_Westerling_et_al__2011.pdf

130. White JW, Hoogenboom G, Kimball BA, Wall 

GW. 2011. Methodologies for simulating impacts of climate 

change on crop production. Field Crops Research 124:357-368.

Type: Journal 

Geographic Area: Global

Keywords: adaptation, agricultural impacts, climate change, 

crop growth simulation, global warming, modeling

Abstract: Ecophysiological models are widely used to forecast potential impacts of climate change on future agricultural 

productivity and to examine options for adaptation by local 

stakeholders and policy makers. However, protocols followed 

in such assessments vary to such an extent that they constrain 

distribution and their contemporary realized climatic niche 

rapidly disassociates through the century. These models demonstrate the heightened risk for species occupying small geographic ranges of displacement into climatic disequilibrium 

from rapid climate change and provide tools to assist decision 

makers in mitigating the threat.

Link: http://www.treesearch.fs.fed.us/pubs/37041

128. Wasserman TN, Cushman SA, Shirk AS, 

Landguth EL, Littell JS. 2011. Simulating the effects of 

climate change on population connectivity of American marten 

(Martes americana) in the northern Rocky Mountains, USA. 

Landscape Ecology 27:211-225.

Type: Journal 

Geographic Area: Northwestern USA

Keywords: habitat fragmentation, gene flow, climate change, 

American marten, Martes americana, connectivity, landscape 

genetics

Abstract: We utilize empirically derived estimates of landscape resistance to assess current landscape connectivity of 

American marten (Martes americana) in the northern Rocky 

Mountains, USA, and project how a warming climate may 

affect landscape resistance and population connectivity in 

the future. We evaluate the influences of five potential future 

temperature scenarios involving different degrees of warming. We use resistant kernel dispersal models to assess population connectivity based on full occupancy of suitable habitat 

in each of these hypothetical future resistance layers. We use 

the CDPOP model to simulate gene exchange among individual martens in each of these hypothetical future climates. 

We evaluate: (1) changes in the extent, connectivity and pattern of marten habitat, (2) changes in allelic richness and expected heterozygosity, and (3) changes in the range of significant positive genetic correlation within the northern Idaho 

marten population under each future scenario. We found that 

even moderate warming scenarios resulted in very large reductions in population connectivity. Calculation of genetic 

correlograms for each scenario indicates that climate driven 

changes in landscape connectivity results in decreasing range 

of genetic correlation, indicating more isolated and smaller 

genetic neighborhoods. These, in turn, resulted in substantial 

loss of allelic richness and reductions in expected heterozygosity. In the U.S. northern Rocky Mountains, climate change 

may extensively fragment marten populations to a degree that 

strongly reduces genetic diversity. Our results demonstrate 

that for species, such as the American marten, whose population connectivity is highly tied to climatic gradients, expected 

climate change can result in profound changes in the extent, 

pattern, connectivity and gene flow of populations.

Link: http://www.fs.fed.us/rm/pubs_other/rmrs_2012_wasserman_t002.pdf

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
46 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Abstract: Vegetation records spanning the past 21 kyr in 

western North America display spatial patterns of change that 

reflect the influence of variations in the large-scale controls of 

climate. Among these controls are millennial-scale variations 

in the seasonal cycle of insolation and the size of the ice sheet, 

which affect regional climates directly through changes in 

temperature and net radiation, and indirectly by shifting atmospheric circulation. Longer vegetation records provide an opportunity to examine the regional response to different combinations of these large-scale controls, and whether non-climatic 

controls are important. But most of the longer North American 

records are of insufficient quality to allow a robust test, and 

the long European records are in regions where the vegetation response to climate is often difficult to separate from the 

response to ecological and anthropogenic controls. Here we 

present a 125-kyr record of vegetation and climate change for 

the forest/steppe border of the eastern Cascade Range, northwest America. Pollen data disclose alternations of forest and 

steppe that are consistent with variations in summer insolation 

and global ice volume, and vegetational transitions correlate 

well with the marine isotope stage boundaries. The close relationship between vegetation and climate beyond the Last 

Glacial Maximum provides evidence that climate variations 

are the primary cause of regional vegetation change on millennial timescales, and that non-climatic controls are secondary.

132. Williams JW, Jackson ST. 2007. Novel climates, 

no-analog communities, and ecological surprises. Frontiers in 

Ecology and the Environment 5:475-482.

Type: Journal 

Geographic Area: North America

Compilers’ Keywords: novel ecosystems, no-analogue plant 

associations, climate change, IPCC, modern analogs, greenhouse emissions

Abstract: No-analog communities (communities that are 

compositionally unlike any found today) occurred frequently in the past and will develop in the greenhouse world of 

the future. The well documented no-analog plant communities of late-glacial North America are closely Linked to 

“novel” climates also lacking modern analogs, characterized 

by high seasonality of temperature. In climate simulations 

for the Intergovernmental Panel on Climate Change A2 and 

B1 emission scenarios, novel climates arise by 2100 AD, 

primarily in tropical and subtropical regions. These future 

novel climates are warmer than any present climates globally, with spatially variable shifts in precipitation, and increase the risk of species reshuffling into future no-analog 

communities and other ecological surprises. Most ecological models are at least partially parameterized from modern 

observations and so may fail to accurately predict ecological 

responses to these novel climates. There is an urgent need 

to test the robustness of ecological models to climate conditions outside modern experience.

cross-study syntheses and increase the potential for bias in 

projected impacts. We reviewed 221 peer reviewed papers 

that used crop simulation models to examine diverse aspects 

of how climate change might affect agricultural systems. 

Six subject areas were examined: target crops and regions; 

the crop model(s) used and their characteristics; sources and 

application of data on CO2 and climate; impact parameters 

evaluated; assessment of variability or risk; and adaptation 

strategies. Wheat, maize, soybean and rice were considered 

in approximately 170 papers. The USA (55 papers) and 

Europe (64 papers) were the dominant regions studied. The 

most frequent approach used to simulate response to CO2

involved adjusting daily radiation use efficiency (RUE) and 

transpiration, precluding consideration of the interacting effects of CO2, stomatal conductance and canopy temperature, 

which are expected to exacerbate effects of global warming. 

The assumed baseline CO2 typically corresponded to conditions 10–30 years earlier than the date the paper was accepted, 

exaggerating the relative impacts of increased CO2. Due in 

part to the diverse scenarios for increases in greenhouse gas 

emissions, assumed future CO2 also varied greatly, further 

complicating comparisons among studies. Papers considering 

adaptation predominantly examined changes in planting dates 

and cultivars; only 20 papers tested different tillage practices 

or crop rotations. Risk was quantified in over half the papers, 

mainly in relation to variability in yield or effects of water 

deficits, but the limited consideration of other factors affecting risk beside climate change per se suggests that impacts 

of climate change were overestimated relative to background 

variability. A coordinated crop, climate and soil data resource 

would allow researchers to focus on underlying science. More 

extensive model intercomparison, facilitated by modular software, should strengthen the biological realism of predictions 

and clarify the limits of our ability to forecast agricultural 

impacts of climate change on crop production and associated 

food security as well as to evaluate potential for adaptation.

Link: http://digitalcommons.unl.edu/cgi/viewcontent.cgi?arti

cle=1891&context=usdaarsfacpub&sei-redir=1&referer=http

%3A%2F%2Fscholar.google.com%2Fscholar%3Fq%3DMeth

odologies%2Bfor%2Bsimulating%2Bimpacts%2Bof%2Bclim

ate%2Bchange%2Bon%2Bcrop%2Bproduction%26btnG%3D

%26hl%3Den%26as_sdt%3D0%252C13#search=%22Method

ologies%20simulating%20impacts%20climate%20change%20

crop%20production%22

131. Whitlock C, Bartlein PJ. 1997. Vegetation and 

climate change in the northwest America during the past 125 kyr. 

Nature 388:57-61.

Type: Journal 

Geographic Area: Western North America

Compilers’ Keywords: migration, pollen record, glacial 

maximum, Cascade Range, past vegetation

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 47

in the no-analogue pollen assemblages, nor can they explain 

the prevalence of no-analogue pollen samples during the late 

glacial period.

Link: http://www.cof.orst.edu/cof/teach/fs545/Readings/

Williams,%20Shuman,%20Webb%202001.pdf

134. Wilsey CB, Lawler JJ, Maurer EP, McKenzie 

D, Townsend PA, Gwozdz R, Freund JA, Hagmann 

K, Hutten KM. 2013. Tools for assessing climate impacts 

on fish and wildlife. Journal of Fish and Wildlife Management 

4:220-241.

Type: Journal 

Geographic Area: North America

Keywords: climate change, ecological modeling, hydrology, 

vegetation, fire

Abstract: Climate change is already affecting many fish and 

wildlife populations. Managing these populations requires an 

understanding of the nature, magnitude, and distribution of 

current and future climate impacts. Scientists and managers 

have at their disposal a wide array of models for projecting 

climate impacts that can be used to build such an understanding. Here, we provide a broad overview of the types of models 

available for forecasting the effects of climate change on key 

processes that affect fish and wildlife habitat (hydrology, fire, 

and vegetation), as well as on individual species distributions 

and populations. We present a framework for how climate-impacts modeling can be used to address management concerns, 

providing examples of model-based assessments of climate 

impacts on salmon populations in the Pacific Northwest, fire 

regimes in the boreal region of Canada, prairies and savannas 

in the Willamette Valley-Puget Sound Trough-Georgia Basin 

ecoregion, and marten populations in the northeastern United 

States and southeastern Canada. We also highlight some key 

limitations of these models and discuss how such limitations 

should be managed. We conclude with a general discussion 

of how these models can be integrated into fish and wildlife 

management.

135. Worrall JJ, Rehfeldt GE, Hamann A, Hogg 

EH, Marchetti SB, Michaelian M, Gray LK. 2013.

Recent declines of Populus tremuloides in North America linked 

to climate. Forest Ecology and Management 299:35-51.

Type: Journal 

Geographic Area: North America

Keywords: decline, dieback, die-off, drought, climate envelope, climatic niche

Abstract: Populus tremuloides (trembling aspen) recently 

experienced extensive crown thinning, branch dieback, and 

mortality across North America. To investigate the role of 

climate, we developed a range wide bioclimate model that 

characterizes climatic factors controlling distribution of aspen. 

Link: http://training.fws.gov/EC/Resources/climate_change/

lcc/nov_10/williams_and_jackson%20_2007_frontiers.pdf

133. Williams JW, Shuman BN, Webb TI. 2001.

Dissimilarity analyses of late-quaternary vegetation and climate 

in eastern North America. Ecology 82:3346-3362.

Type: Journal 

Geographic Area: Eastern North America

Keywords: biomes, dissimilarity analysis, eastern North 

America, equilibrium vs. disequilibrium, hypotheses, noanalogue plant associations, paleoecology, paleoclimatology, 

pollen, Quaternary, vegetation

Abstract: Plant formations different from any extant today 

apparently were widespread in North America and Europe 

during the last deglaciation, produced by the independent 

biogeographic responses of plant taxa to climate change. 

Dissimilarity analyses of modern and fossil pollen samples 

in eastern North America show that the unique plant associations centered around the Great Lakes at 14,000 calendar 

years before present (yr BP), with high dissimilarities during 

17,000–12,000 yr BP. The late-glacial fossil pollen assemblages are characterized by (1) high abundances of boreal 

conifers such as spruce and larch relative to their Holocene 

values, (2) high abundances of herbaceous types (sedge, sage, 

and ragweed), (3) high abundances of broad-leaved deciduous 

types (ash, hornbeam, poplar, hazel, and willow), and (4) the 

low abundance or absence of pine, alder, and birch. When the 

fossil pollen samples are assigned to biomes using the affinity core technique, the late Pleistocene pollen samples are assigned to mixed parkland, a biome that is not extant in North 

America today. The fastest vegetational changes occurred 

13,000–11,000 yr BP, when the late Pleistocene vegetation reorganized into the Holocene biomes, which have persisted to 

today. Simulations by the Community Climate Model, version 

1 (CCM1), suggest that late glacial climates were also unlike 

modern climates, featuring a ‘‘hypercontinental’’ mixture of 

colder-than-present winters, warmer than-present summers, 

and lower-than-present precipitation. Dissimilarity analyses 

of the pollen data and CCM1 simulations for 21,000, 16,000, 

14,000, 11,000, and 6,000 yr BP show that (1) the temporal 

and spatial distribution of high dissimilarities in the vegetation 

(relative to present) coincide with dissimilarities in simulated 

climate, (2) the timing and spatial distribution of changes in the 

vegetation and simulated climate also agree, and (3) the largest climatic and vegetational changes follow the peak period 

of dissimilarity from present. Taken together, these three lines 

of evidence support the hypothesis that the no-analogue plant 

associations were in equilibrium with orbital- and millennialscale climate change. Nonclimatic factors such as low atmospheric CO2 concentrations and the presence of now-extinct 

megafauna species may have increased the openness of the 

Pleistocene vegetation, but by themselves cannot explain the 

observed mixture of boreal, temperate, and herbaceous taxa 

CHAPTER 1 • CLIMATE CHANGE • RESEARCH
48 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

We also examined indices of moisture stress, insect defoliation 

and other factors as potential causes of the decline. Historic 

climate records show that most decline regions experienced 

exceptionally severe drought preceding the recent episodes. 

The bioclimate model, driven primarily by maximum summer 

temperatures and April–September precipitation, shows that 

decline tended to occur in marginally suitable habitat, and that 

climatic suitability decreased markedly in the period leading 

up to decline in almost all decline regions. Other factors, notably multi-year defoliation by tent caterpillars (Malacosoma

spp.) and stem damage by fungi and insects, also play a substantial role in decline episodes, and may amplify or prolong 

the impacts of moisture stress on aspen over large areas. Many 

severely affected stands have poor regeneration potential, raising concerns that increasing aridity could ultimately lead to 

widespread loss of aspen forest cover. The analysis indicates 

that exceptional droughts were a major cause of the decline 

episodes, especially in the drier regions, and that aspen is sensitive to drought in much of its range. Coupling the bioclimate model with climate projections suggests that we should 

expect substantial loss of suitable habitat within the current 

distribution, especially in the USA and Mexico.

Link: http://www.ualberta.ca/~ahamann/publications/pdfs/

Worrall_et_al_2013.pdf
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 49

136. Alfaro RI, Fady B, Vendramin GG, Dawson 

IK, Fleming RA, Sáenz-Romero C, Lindig-Cisneros 

RA, Murdock T, Vinceti B, Navarro CM, Skrøppa 

T, Baldinelli G, El-Kassaby YA, Loo J. 2014. The role 

of forest genetic resources in responding to biotic and abiotic 

factors in the context of anthropogenic climate change. Forest 

Ecology and Management 333:76-87.

Type: Journal 

Geographic Area: Global

Keywords: tree genetic variation, climate change, adaptation, 

natural disturbance

Abstract: The current distribution of forest genetic resources 

on Earth is the result of a combination of natural processes 

and human actions. Over time, tree populations have become 

adapted to their habitats including the local ecological disturbances they face. As the planet enters a phase of human-induced climate change of unprecedented speed and magnitude, 

however, previously locally-adapted populations are rendered 

less suitable for new conditions, and “natural” biotic and abiotic disturbances are taken outside their historic distribution, 

frequency and intensity ranges. Tree populations rely on phenotypic plasticity to survive in extant locations, on genetic 

adaptation to modify their local phenotypic optimum or on migration to new suitable environmental conditions. The rate of 

required change, however, may outpace the ability to respond, 

and tree species and populations may become locally extinct 

after specific, but as yet unknown and unquantified, tipping 

points are reached. Here, we review the importance of forest 

genetic resources as a source of evolutionary potential for adaptation to changes in climate and other ecological factors. We 

particularly consider climate-related responses in the context 

of Linkages to disturbances such as pests, diseases and fire, 

and associated feedback loops. The importance of management strategies to conserve evolutionary potential is emphasised and recommendations for policy-makers are provided.

Link: http://www.sciencedirect.com/science/article/pii/

S037811271400231X

137. Anderson PD, Chmura DJ. 2009. Silvicultural 

approaches to maintain forest health and productivity under 

current and future climates. Western Forester 54:6-8.

Type: Newsletter

Geographic Area: Pacific Northwest USA

Compilers’ Keywords: climate change, forest management

Introduction: Climate modeling based on a variety of scenarios for the Pacific Northwest suggests that over the next 

century temperatures may increase and that the abundance 

of summer precipitation may decline. Historically, climate 

General (1–36)—editorials, opinions, 

reviews, trends, surveys, summaries

Research (37–135)—climate change impacts, 

projections, paleoecology

Strategies (136–205)—policies, adaptation 

options, frameworks, mitigations

Resources (206–214)—tools, websites, software

Climate Change 

CHAPTER 1 • CLIMATE CHANGE • STRATEGIES

#. Author(s), Date. Title. Source.

Type: 

Geographic Area: 

Keywords: 

Description:

Link:

location of study

keywords written by the 

author(s) or provided by the 

compilers

ID number and

information

abstract, summary, or 

portions of the introduction 

or conclusion written by the 

author(s) or provided by the 

compilers

book, book section, conference 

paper, dissertation, government 

document, journal article, 

newsletter, presentation, report, 

thesis or web page

internet address of reference

Each reference contains the following:
50 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

of climate change for the provision of such ecosystem services 

at landscape levels.

Link: http://curis.ku.dk/ws/files/20572936/wp31.pdf

139. Campbell EM, Saunders SC, Coates KD, 

Meidinger DV, MacKinnon A, O’Neill GA, 

MacKillop DJ, DeLong SC, Morgan DG. 2009.

Ecological resilience and complexity: a theoretical framework for 

understanding and managing British Columbia’s forest ecosystem in a changing climate. Victoria, British Columbia, Canada: 

BC Ministry of Forests and Range, Forest Science Program. 

Technical Report 055. 44 p.

Type: Government Document

Geographic Area: Western Canada

Compilers’ Keywords: climate change, forest management, 

mitigation, ecosystem services, review

Summary: At global, regional, and local scales, forest managers are faced with unprecedented pressures to supply forest 

resources for human consumption while still maintaining a 

diverse array of other ecosystem services essential to human 

well-being. While this alone has posed major challenges to forest management, global climate change presents a new range of 

daunting challenges. The potential for major ecosystem changes as well as uncertainties about the degree and rate of climate 

change necessitates a major shift in thinking about forest management. Recent scientific literature proposes new approaches 

to forest management that focus on “managing for ecological 

resilience,” with the idea that it provides a tenable framework 

for achieving sustainability goals when environments are 

changing and the future is uncertain. The concept of ecological 

resilience has been used to guide the management of ecosystems degraded by human land use activities, and managing for 

resilience is a commonly discussed approach for countering 

the negative impacts of climate change. This document summarizes the theoretical literature on ecological resilience and 

complexity, and describes how this evolving body of science 

can begin to guide the management of forest ecosystems in a 

changing climate. Ecological resilience describes the capacity 

of ecosystems to absorb disturbance without collapsing into a 

qualitatively different state. While research is still under way to 

develop a structured understanding of the mechanisms regulating ecological resilience, scientists propose that key ecological 

processes, operating across varying scales of time and space 

(e.g., seedling survivorship, forest succession, periodic natural 

disturbances, propagule dispersal that facilitates species range 

shifts), generate the complexity needed to maintain ecosystem 

resilience to environmental change. Maintaining and enhancing biological diversity across multiple scales may play an 

important role in preserving ecosystem services if it generates 

redundancy in the ecological processes that confer ecological 

resilience (i.e., conserves key ecosystem functions). Adapting 

forest management frameworks to climate change involves 

changes at the century scale have been accompanied by adjustments in species population sizes and the composition of vegetation communities. Silvicultural practices may be employed 

to assist in forest adaptation to climate changes, principally by 

influencing stand structure and species composition throughout stand development. Forest managers might consider a primary silviculture objective over the next century of increasing 

forest vigor to increase both resistance and resilience to direct 

and indirect stresses arising from climate variation.

Link: http://www.fs.fed.us/pnw/pubs/journals/pnw_2009_anderson001.pdf

138. Bosselmann AS, Jacobsen JB, Kjaer ED, 

Thorsen BJ. 2008. Climate change, uncertainty and the economic value of genetic diversity: a pilot study on methodologies. 

Forest and Landscape Working Papers 31-2008:1-58.

Type: Journal 

Geographic Area: Denmark, Europe

Compilers’ Keywords: methods, forest management

Summary: The report is based on a limited empirical material, which stresses the need for qualified information from 

genetic research and tree improvement programmes as climate 

change is picking up speed and forest owners every day make 

decisions that will affect the stability, health and growth of 

forest many decades into the future. The decisions are made 

on information that was produced under a research and tree 

improvement paradigm focused on tree breeding for production under a known climate believed to be stable. Any new 

and improved information is greatly needed—even if it is not 

exact and only indicative. Therefore, existing clone and provenance trials in forest genetic research and tree improvement 

programmes should be systematically investigated across as 

large climate gradients as permitted by the location of the trials. Such analysis should be able to bring about at least indicative information on the genotype response to variation in the 

main climate and growth factors. Such information can subsequently be used in more thorough analyses along the steps 

outlined here, perhaps with a special focus on potential gains 

from mixing provenances on the same area. Tree improvement 

programmes should use the information produced to gradually 

adjust current recommendations concerning local and regional 

choices of planting material, and of course to adjust and diversify breeding strategies accordingly. In the long run, such 

research may go into more details along the lines discussed 

earlier. In particular, genetic research into sensitivity of different provenances and clones to potentially more frequent 

pest attacks, mild winters, etc. may be relevant. On a broader 

scale, ecological and economic research could aim to provide 

more information on the way forest ecosystem services and 

their value rely on the state and characteristics of the forest 

ecosystem and notably its stability. Such information will be 

needed for a reliable economic modelling of the consequences 

CHAPTER 1 • CLIMATE CHANGE • STRATEGIES
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 51

Keywords: ecosystem services, forest restoration, bioenergy, 

climate change, planting stock production, economics

Abstract: Global deforestation and forest degradation have 

led to massive loss of biodiversity and decline of ecosystem 

services. Against this prospect, it is important not only to protect, but also to restore forest ecosystems. The paper analyzes 

the current and future role of the restoration of forests and degraded lands starting with the definition of various techniques, 

scales and objectives of forest restoration. Three key motivations for and targets of forest restoration are then discussed: 

forest biodiversity protection, biomass production, climate 

change mitigation and adaptation. The paper also briefly 

discusses three tools of increasing relevance for supporting 

forest restoration policies: the development of forest nursery 

techniques and the improvement of quality of forest reproductive material, the use of standard and certification tools, and 

financing of restoration investments, including projects aimed 

at reducing emissions from deforestation and forest degradation. We conclude by making some final remarks on the future 

challenges of forest restoration policies. 

141. Colombo S, Boysen B, Brosemer K, Foley A, 

Obenchain A. 2008. Managing tree seed in an uncertain climate: conference summary. Ontario, Canada: Ontario Ministry 

of Natural Resources, Applied Research and Development 

Branch. Note Number 8. 8 p.

Type: Government Document

Geographic Area: Central Canada

Compilers’ Keywords: forest, seed transfer, adaptive variation, forest genetic resources, forest management

Introduction: Canadian forests will experience greater 

increases in temperature than those growing further south. 

One set of projections for Ontario indicates that by the end of 

the century, average summer temperature will increase from 

3 to 7 °C (southern and northern Ontario, respectively) and 

that winter temperatures will increase from up to 4 °C in the 

south to 10 °C in the north. Predictions of precipitation with 

climate change vary, but future precipitation may be insufficient to prevent large areas of Ontario forests from experiencing more frequent and intense drought. Such temperature 

and soil moisture changes would alter the ecological conditions in forests enough to affect how well trees are adapted 

to local climates. As a result, climate change presents many 

challenges to sustainable forest management in Canada. The 

long life of tree species and long rotation cycles mean that the 

effects of today’s decisions can persist for a century or more. 

Given projections for Ontario’s future climate, those who are 

concerned about forest ecology, management, and policy must 

be concerned about seed source deployment. Important questions include: 1. What if any changes do we make in how we 

distribute seed sources when planting? 2. When do we make 

changes? 3. Can we predict the effects of such changes? Out of 

actions that minimize the risk of adverse climate-change impacts and capitalize on its benefits. Managing for resilience 

advocates diverse and novel actions that help cope with uncertainties about future forest conditions and reduce both societal and ecological vulnerabilities to climate change. While 

societal adaptations to climate change include the development 

of policies to encourage adaptation, modifying wood processing technologies, and revising expectations of resource use and 

conservation objectives, management to maintain ecological 

resilience involves deliberate, on-the-ground forest practices 

that maintain ecosystem complexity across multiple scales of 

time and space, and facilitate gradual ecosystem change in 

response to climate change. In this report, we present some 

examples of the kinds of on-the-ground actions that could be 

undertaken to begin the process of managing for ecological 

resilience to climate change. They include: facilitating tree 

species (and population) migration and range shifts; developing forest harvest, regeneration, and stand-tending activities 

that maintain or enhance ecosystem complexity and response 

diversity to environmental change, such as the forest structures generated by past disturbance regimes; planting broader 

mixes of trees across landscapes to help reset successional trajectories; promoting landscape connectivity; and retaining or 

restoring areas that may be buffered against climate change. 

Managing for ecological resilience is in its infancy and the 

technical details about how to implement this new approach to 

forest management will depend on ecosystem type and evolve 

as the science integrating resilience, complexity, and biodiversity evolves and more information about the impacts of climate change on ecosystems becomes available through field 

monitoring programs and quantitative modelling research. 

Resilience-based ecosystem management, especially when it 

takes into account changing climate conditions, represents a 

profound shift in the way the Ministry of Forests and Range 

will approach how forest ecosystems are managed, and poses 

challenges to many existing practices and policies in British 

Columbia. Involving a more diverse array of forest practices 

than traditional forest management approaches, it will require 

setting landscape-level management objectives for desired future forest conditions and making decisions regarding how to 

cope with unexpected and undesirable management outcomes. 

Transitioning to resilience-based ecosystem management also 

requires an understanding of complex ecological and social 

feedbacks, but an awareness of this, and an openness to work 

with complexity scientists, forest ecologists, climatologists, 

forest geneticists, and others can help to make this transition.

Link: http://www.for.gov.bc.ca/hfd/pubs/Docs/Tr/Tr055.htm

140. Ciccarese L, Mattsson A, Pettenella D. 2012.

Ecosystem services from forest restoration: thinking ahead. New 

Forests 43:543-560.

Type: Journal 

Geographic Area: Global

CHAPTER 1 • CLIMATE CHANGE • STRATEGIES
52 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

143. Driscoll DA, Felton A, Gibbons P, Felton AM, 

Munro NT, Lindenmayer DB. 2011. Priorities in policy 

and management when existing biodiversity stressors interact 

with climate-change. Climatic Change 111:533-557.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: environmental stressors

Abstract: There are three key drivers of the biodiversity crisis: 1) the well-known existing threats to biodiversity such as 

habitat loss, invasive pest species and resource exploitation; 2) 

direct effects of climate-change, such as on coastal and high 

elevation communities and coral reefs; and 3) the interaction 

between existing threats and climate-change. The third driver 

is set to accelerate the biodiversity crisis beyond the impacts of 

the first and second drivers in isolation. In this review we assess these interactions, and suggest the policy and management 

responses that are needed to minimise their impacts. Renewed 

management policy action that address known threats to biodiversity could substantially diminish the impacts of future 

climate change. An appropriate response to climate-change 

will include a reduction of land clearing, increased habitat restoration using indigenous species, a reduction in the number 

of exotic species transported between continents or between 

major regions of endemism, and a reduction in the unsustainable use of natural resources. Achieving these measures requires substantial reform of international, national and regional 

policy, and the development of new or more effective alliances 

between scientists, government agencies, non-government 

organisations and land managers. Furthermore, new management practices and policy are needed that consider shifts in the 

geographic range of species, and that are responsive to new 

information acquired from improved research and monitoring 

programs. The interactions of climate-change with existing 

threats to biodiversity have the potential to drive many species 

to extinction, but there is much that can be done now to reduce 

this risk.

144. Erickson VJ, Aubry C, Berrang PC, Blush 

T, Bower AD, Crane BS, DeSpain T, Gwaze D, 

Hamlin J, Horning ME, Johnson R, Mahalovich 

MF, Maldonado M, Sniezko R, St. Clair JB. 2012.

Genetic resource management and climate change: genetic 

options for adapting national forests to climate change. 

Washington (DC): USDA Forest Service, Forest Management. 

24 p.

Type: Government Document

Geographic Area: North America

Compilers’ Keywords: genecology, planning, USFS, genetic 

options, review, guidelines

Summary: This report provides an overview of current climate change knowledge and potential implications for forest 

concern for these issues, a conference entitled Managing Tree 

Seed in an Uncertain Climate was held on November 14-15, 

2007, in Sault Ste. Marie, Ontario. The event was co-hosted by 

the Ontario Ministry of Natural Resources (through its climate 

change initiative) and Forest Genetics Ontario. This conference focused on managing tree seed in a rapidly changing climate, including seed movement (species and populations) and 

regulatory tools (guidelines/zones); the science behind identifying and predicting patterns of adaptive variation response to 

climate change (invited speakers’ abstracts are shown in the 

appendix); the need to revise seed movement decision-making; and reforestation aspects of management planning under 

climate change. The goals of the conference were to: 1. Build 

awareness about climate change and its potential impacts on 

the deployment of forest genetic resources (e.g., seed transfer guidelines, seed source bulking, seed banking, and tree 

improvement strategies) 2. Initiate a dialogue among those 

concerned about impacts of climate change on forest genetic 

resources 3. Develop a roadmap of future actions promoting 

adaptation of forests to future climate.

Link: http://www.mnr.gov.on.ca/stdprodconsume/groups/

lr/@mnr/@climatechange/documents/document/276918.pdf

142. Conroy MJ, Runge MC, Nichols JD, Stodola 

KW, Cooper RJ. 2011. Conservation in the face of climate 

change: the roles of alternative models, monitoring, and adaptation in confronting and reducing uncertainty. Biological 

Conservation 144:1204-1213.

Type: Journal 

Geographic Area: Eastern USA

Keywords: climate change, birds, prediction, monitoring, 

adaptive management

Abstract: The broad physical and biological principles behind 

climate change and its potential large scale ecological impacts 

on biota are fairly well understood, although likely responses 

of biotic communities at fine spatio-temporal scales are not, 

limiting the ability of conservation programs to respond effectively to climate change outside the range of human experience. 

Much of the climate debate has focused on attempts to resolve 

key uncertainties in a hypothesis-testing framework. However, 

conservation decisions cannot await resolution of these scientific issues and instead must proceed in the face of uncertainty. 

We suggest that conservation should precede in an adaptive 

management framework, in which decisions are guided by 

predictions under multiple, plausible hypotheses about climate 

impacts. Under this plan, monitoring is used to evaluate the 

response of the system to climate drivers, and management 

actions (perhaps experimental) are used to confront testable 

predictions with data, in turn providing feedback for future decision making. We illustrate these principles with the problem 

of mitigating the effects of climate change on terrestrial bird 

communities in the southern Appalachian Mountains, USA.

CHAPTER 1 • CLIMATE CHANGE • STRATEGIES
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 53

for those who are aware of them, practical guidance on how to 

respond is largely non-existent. The extent to which forestry 

research and national policies will promote and adopt management practices in order to assist production forests adapt to climate change is currently uncertain. Mainstreaming adaptation 

into national development and planning programs may represent an initial step towards the incorporation of climate change 

considerations into tropical forestry.

Link: http://www.cifor.org/publications/pdf_files/articles/

AGuariguata0801.pdf

146. Gunn JS, Hagan JM, Whitman AA. 2009. 

Forestry adaptation and mitigation in a changing climate—a forest resource manager’s guide for the northeastern United States. 

Brunswick (ME): Natural Capital Initiative at Manomet. NCI2009-1. 16 p.

Type: Report

Geographic Area: Northeastern USA

Compilers’ Keywords: climate change, forest management, 

guidelines, silviculture, report, resilience

Summary: A coordinated response to reduce greenhouse 

gas emissions will not be adequate to prevent unprecedented 

climate change resulting from the elevated levels of carbon 

dioxide (CO2) already in the atmosphere. Climate adaptation 

management strategies are essential if landowners and managers are going to have plans that help them achieve their objectives in the face of uncertainty (e.g., revenue, silviculture, retain third-party forest certification). Strategies are also needed 

if northeastern U.S. forests will continue to play a mitigating 

role in addressing greenhouse gas emissions. In this report, we 

lay out a framework for understanding potential impacts of climate change on forestry. This framework draws on a review 

of recommended actions from forest managers and scientists 

throughout Canada, the U.S., and Europe. We then present a 

toolbox of practices that forest managers in the northeastern 

U.S. might apply to reduce exposure to the immediate and 

long-term risk from climate change. The toolbox approach 

incorporates three broad strategies of Resistance, Resilience, 

and Response. A Resistance strategy is a set of short-term approaches to address immediate threats and focuses on minimizing the impacts of disturbance regimes that are exacerbated 

by climate change. Resilience can be seen both as a short-term 

and a long-term strategy. Resilience strategies address the capacity of a stand or community to recover from a disturbance 

and return to a reference or desired state. The primary purpose 

of a Response strategy is to facilitate the movement of species 

over time. This strategy encompasses the most costly practices 

and requires acceptance of a level of uncertainty that many 

landowners and managers will likely not choose.

Link: http://www.manomet.org/sites/default/files/publications_and_tools/Manomet_ForestryAdaptationtoCCReport%205-09.pdf

tree species, as well as goals, principles, and recommendations for enhancing forest resilience and resistance through 

a re-aligned “climate-smart” National Forest System (NFS) 

Genetic Resource Management Program. Although national 

forests may differ in terms of species and population vulnerability to climate change, as well as appropriate management 

response, our recommendations and adaptation options all follow three overarching principles: (1) genetically diverse and 

adapted seed and planting stock will provide the foundation for 

healthy forests and ecosystems in the future; (2) gene conservation is key to preserving vulnerable species and populations 

for the future; and (3) establishing and maintaining partnerships will be more important than ever. Implementation of the 

adaptation options will require new tools, practices, and refocused investments in NFS Genetic Resource Management 

activities, as well as a trained workforce, supporting plant production infrastructure, and strong support from research and 

management.

Link: http://www.fs.usda.gov/Internet/FSE_DOCUMENTS/

stelprdb5368468.pdf

145. Guariguata MR, Cornelius JP, Locatelli B, 

Forner C, Sánchez-Azofeifa GA. 2008. Mitigation 

needs adaptation: tropical forestry and climate change.

Mitigation and Adaptation Strategies for Global Change 

13:793-808.

Type: Journal 

Geographic Area: Global 

Keywords: climate change, adaptation, tropical forests, tropical tree plantations, natural forest management

Abstract: The relationship between tropical forests and global climate change has so far focused on mitigation, while much 

less emphasis has been placed on how management activities 

may help forest ecosystems adapt to this change. This paper 

discusses how tropical forestry practices can contribute to 

maintaining or enhancing the adaptive capacity of natural and 

planted forests to global climate change and considers challenges and opportunities for the integration of tropical forest 

management in broader climate change adaptation. In addition 

to the use of reduced impact logging to maintain ecosystem 

integrity, other approaches may be needed, such as fire prevention and management, as well as specific silvicultural options 

aimed at facilitating genetic adaptation. In the case of planted 

forests, the normally higher intensity of management (with 

respect to natural forest) offers additional opportunities for implementing adaptation measures, at both industrial and smallholder levels. Although the integration in forest management 

of measures aimed at enhancing adaptation to climate change 

may not involve substantial additional effort with respect to 

current practice, little action appears to have been taken to 

date. Tropical foresters and forest-dependent communities appear not to appreciate the risks posed by climate change and, 

CHAPTER 1 • CLIMATE CHANGE • STRATEGIES
54 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

these challenges. Five elements of CCS are described: regional 

modelling, expanding protected areas; management of the matrix regional coordination; and transfer of resources. Regional 

modelling uses regional climate models, biotic response models and sensitivity analysis to identify climate change impacts 

on biodiversity at a regional scale appropriate for conservation planning. Expansion of protected areas management and 

systems within the planning region are based on modelling 

results. Management of the matrix between protected areas 

provides continuity for processes and species range shifts 

outside of parks. Regional coordination of park and off-park 

efforts allows harmonization of conservation goals across 

provincial and national boundaries. Finally, implementation 

of these CCS elements in the most biodiverse regions of the 

world will require technical and financial transfer of resources 

on a global scale. Collaboration across disciplines is necessary 

to plan conservation responses to climate change adequately. 

Biogeography and ecology provide insights into the effects 

of climate change on biodiversity that have not yet been fully 

integrated into conservation biology and applied conservation 

management. CCS provide a framework in which biogeographers, ecologists and conservation managers can collaborate 

to address this need. These planning exercises take place on 

a regional level, driven by regional climate models as well as 

general circulation models (GCMs), to ensure that regional 

climate drivers such as land use change and mesoscale topography are adequately represented. Sensitivity analysis can help 

address the substantial uncertainty inherent in projecting future 

climates and biodiversity response.

Link: http://www.cakex.org/sites/default/files/Climate%20

change%20integrated%20conservation%20strategies_1.pdf

149. Hayward, GD, Flather, CH, Uloth, E, Safford, 

HD, Cleaves, DA. 2009. Managing fish and wildlife in the 

face of climate change: USDA Forest Service perspective. In: 

McCabe RE, Stockwell KA, editors. Transactions of the 74th 

North American Wildlife and Natural Resources Conference. 

Washington (DC): Wildlife Management Institute. p 98-109. 

Type: Proceedings Paper

Geographic Area: North America

Compilers’ Keywords: climate change, assisted migration, 

fish and wildlife management, IPCC, strategic framework, review, USFS

Introduction: This invited review explores progress that the 

research and management branches of the Forest Service have 

made addressing fish and wildlife management in the face of 

climate change. In this paper, we begin by providing an overview of the agency’s research and management context. Key 

to this overview is the Forest Service Strategic Framework 

(U.S. Forest Service 2008a) for responding to climate change, 

which defines three broad categories of agency goals that 

guide the incorporation of climate change into natural resource 

147. Halofsky JE, Peterson DL, Furniss MJ, Joyce 

LA, Millar CI, Neilson RP. 2011. Workshop approach 

for developing climate change adaptation strategies and actions 

for natural resource management agencies in the United States. 

Journal of Forestry 109:219-225.

Type: Journal 

Geographic Area: USA

Keywords: climate change, adaptation, forest management

Abstract: Concrete ways to adapt to climate change are needed to help land-management agencies take steps to incorporate 

climate change into management and take advantage of opportunities to balance the negative effects of climate change. 

Because the development of adaptation tools and strategies is 

at an early stage, it is important that ideas and strategies are 

disseminated quickly to advance thinking and practice. Here, 

we offer an example of a successful workshop, focused on 

National Forests in the United States, which allowed quick 

dissemination of ideas and strategies for climate change adaptation in resource management through an interaction between 

scientists and managers. We share both the process used in the 

workshop and the outcome of facilitated dialogue at the workshop. By presenting concrete adaptation methods and showing 

the value of a focused scientist—manager dialogue, we hope 

to motivate the U.S. Forest Service and other natural resources 

agencies to emulate our approach and begin the process of 

adapting to climate change. 

Link: http://www.treesearch.fs.fed.us/pubs/39144

148. Hannah L, Midgley GF, Miller G. 2002. 

Climate change-integrated conservation strategies. Global 

Ecology and Biogeography 11:485-495.

Type: Journal 

Geographic Area: Global

Keywords: biodiversity, climate change, conservation, matrix 

management, modelling, protected areas, range shifts

Abstract: Conservation strategies currently include little consideration of climate change. Insights about the biotic impacts 

of climate change from biogeography and palaeoecology, 

therefore, have the potential to provide significant improvements in the effectiveness of conservation planning. We suggest a collaboration involving biogeography, ecology and applied conservation. The resulting Climate Change-integrated 

Conservation Strategies (CCS) apply available tools to respond 

to the conservation challenges posed by climate change. The 

focus of this analysis is global, with special reference to high 

biodiversity areas vulnerable to climate change, particularly 

tropical montane settings. Current tools from climatology, biogeography and ecology applicable to conservation planning 

in response to climate change are reviewed. Conservation 

challenges posed by climate change are summarized. CCS 

elements are elaborated that use available tools to respond to 

CHAPTER 1 • CLIMATE CHANGE • STRATEGIES
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 55

showing excessive autumn branchlet drop and top die-back, 

likely as a result of increased summer moisture deficits.

Link: http://www.treesearch.fs.fed.us/pubs/31835

151. Heller NE, Zavaleta ES. 2009. Biodiversity management in the face of climate change: a review of 22 years of recommendations. Biological Conservation 142:14-32.

Type: Journal 

Geographic Area: Global

Keywords: conservation, adaptation, reserve planning, landscape connectivity, resilience, global warming

Abstract: Climate change creates new challenges for biodiversity conservation. Species ranges and ecological dynamics 

are already responding to recent climate shifts, and current 

reserves will not continue to support all species they were 

designed to protect. These problems are exacerbated by other 

global changes. Scholarly articles recommending measures to 

adapt conservation to climate change have proliferated over 

the last 22 years. We systematically reviewed this literature to 

explore what potential solutions it has identified and what consensus and direction it provides to cope with climate change. 

Several consistent recommendations emerge for action at 

diverse spatial scales, requiring leadership by diverse actors. 

Broadly, adaptation requires improved regional institutional 

coordination, expanded spatial and temporal perspective, incorporation of climate change scenarios into all planning and 

action, and greater effort to address multiple threats and global 

change drivers simultaneously in ways that are responsive to 

and inclusive of human communities. However, in the case of 

many recommendations the how, by whom, and under what 

conditions they can be implemented is not specified. We synthesize recommendations with respect to three likely conservation pathways: regional planning; site-scale management; 

and modification of existing conservation plans. We identify 

major gaps, including the need for (1) more specific, operational examples of adaptation principles that are consistent 

with unavoidable uncertainty about the future; (2) a practical 

adaptation planning process to guide selection and integration 

of recommendations into existing policies and programs; and 

(3) greater integration of social science into an endeavor that, 

although dominated by ecology, increasingly recommends extension beyond reserves and into human occupied landscapes.

Link: http://people.ucsc.edu/~zavaleta/pubs/Heller%20

and%20Zavaleta%202009_BiolCons09_climate.pdf

152. Jackson ST, Sax DF. 2010. Balancing biodiversity in a changing environment: extinction debt, immigration 

credit and species turnover. Trends in Ecology and Evolution 

25:153-160.

Type: Journal 

Geographic Area: Global

management. Through a series of case studies we demonstrate 

progress the agency has made in each of those broad goal categories. We conclude by reviewing those features of the Forest 

Service that position it as an important partner in addressing 

climate change, and by highlighting some of the management, 

research, and policy opportunities that must be seized if fish 

and wildlife are to be successfully managed in the face of climate change.

Link: http://www.fs.fed.us/rm/human-dimensions/staff/

flather/pubs/2009.Hayward.et.al.Trans.N.Am.Wildl.Nat.Res.

Conf.pdf

150. Hebda RJ. 2008. Climate change, forests, and the 

forest nursery industry. In: Dumroese RK, Riley LE, editors. National Proceedings: Forest and Conservation Nursery 

Associations – 2007. Fort Collins (CO): USDA Forest Service, 

Rocky Mountain Research Station. Proceedings RMRS-P-57. 

p 81-82.

Type: Government Document

Geographic Area: Western North America

Keywords: ancient carbon, living carbon, mountain pine 

beetle, Dothistroma needle blight

Introduction: The devastating consequences of Hurricane 

Katrina demonstrate how ill-prepared people are when it 

comes to extreme weather events and potential changes in 

climate. The hurricane itself cannot be directly ascribed to climate change, but the likelihood of stronger hurricanes can be. 

The more energy the atmosphere has as it warms because of increasing concentrations of greenhouse gasses, the more energy 

it needs to shuffle around. Hurricanes are one way of doing 

just that. The potential risks from just such an event had been 

described in the region’s major daily paper, yet the response to 

the hurricane seems to indicate that little action had been taken 

to get ready. The lessons of the event must be taken seriously 

by all sectors of society because climate change is a certainty, 

is now well underway, and will impact us all. The fourth series 

of reports issued in 2007 by the Intergovernmental Panel on 

Climate Change (IPCC IV), in what is a conservative account 

of climate change and its impacts, warns us clearly that major effects on forests must be expected. In northwestern North 

America, the climate has already changed and is continuing 

to change, and those changes are having serious impacts on 

regional forests. Furthermore, in one of the IPCC IV reports, 

Fischlin and others (2007) identify northwest North American 

forests as especially likely to be impacted by climate change. 

The devastating mountain pine beetle (Dendroctonus ponderosae) outbreak in the interior of British Columbia and adjacent 

regions has single-handedly altered the character of forests 

over a huge area in less than decade. Increases in Dothistroma 

needle blight on lodgepole pines (Pinus contorta) are also attributed to changes in climate. In the coastal temperate rainforests of British Columbia, western redcedars (Thuja plicata) are 

CHAPTER 1 • CLIMATE CHANGE • STRATEGIES
56 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

or species, innovative pest management, modified enterprise 

structures, and geographic relocation. Social-ecological 

systems in which adaptation is insufficient to counter the 

adverse consequences of climate change might undergo 

transformative change to produce alternative ecosystem services, production enterprises, and livelihoods. The rangeland 

profession is in a pivotal position to provide leadership on 

this global challenge because it represents the intersection 

of management and scientific knowledge, includes diverse 

stakeholders who derive their livelihoods from rangelands, 

and interacts with organizations responsible for rangeland 

stewardship.

Link: http://www.treesearch.fs.fed.us/pubs/44831

154. Joyce LA, Blate GM, McNulty SG, Millar CI, 

Moser S, Neilson RP, Peterson DL. 2009. Managing 

for multiple resources under climate change: national forests. 

Environmental Management 44:1022-1032.

Type: Journal 

Geographic Area: USA

Keywords: resilience, resistance, anticipatory management, 

planning, assessments, adaptation 

Abstract: This study explores potential adaptation approaches in planning and management that the United States 

Forest Service might adopt to help achieve its goals and 

objectives in the face of climate change. Availability of information, vulnerability of ecological and socio-economic 

systems, and uncertainties associated with climate change, 

as well as the interacting non-climatic changes, influence 

selection of the adaptation approach. Resource assessments 

are opportunities to develop strategic information that could 

be used to identify and link adaptation strategies across 

planning levels. Within a National Forest, planning must 

incorporate the opportunity to identify vulnerabilities to climate change as well as incorporate approaches that allow 

management adjustments as the effects of climate change 

become apparent. The nature of environmental variability, 

the inevitability of novelty and surprise, and the range of 

management objectives and situations across the National 

Forest System implies that no single approach will fit all 

situations. A toolbox of management options would include 

practices focused on forestalling climate change effects by 

building resistance and resilience into current ecosystems, 

and on managing for change by enabling plants, animals, 

and ecosystems to adapt to climate change. Better and more 

widespread implementation of already known practices that 

reduce the impact of existing stressors represents an important “no regrets” strategy. These management opportunities 

will require agency consideration of its adaptive capacity, 

and ways to overcome potential barriers to these adaptation 

options.

Link: http://www.ncbi.nlm.nih.gov/pubmed/19588192

Compilers’ Keywords: animals, climate change, humans, 

population dynamics, uncertainty, migration, framework 

Abstract: Here, we outline a conceptual framework for biodiversity dynamics following environmental change. The 

model incorporates lags in extinction and immigration, which 

lead to extinction debt and immigration credit, respectively. 

Collectively, these concepts enable a balanced consideration of 

changes in biodiversity following climate change, habitat fragmentation and other forcing events. They also reveal transient 

phenomena, such as biodiversity surpluses and deficits, which 

have important ramifications for biological conservation and 

the preservation of ecosystem services. Predicting such transient dynamics poses a serious conservation challenge in a 

time of rapid environmental change.

Link: http://www.ncbi.nlm.nih.gov/pubmed/19879014

153. Joyce LA, Briske DD, Brown JR, Polley HW, 

McCarl BA, Bailey DW. 2013. Climate change and North 

American rangelands: assessment of mitigation and adaptation 

strategies. Rangeland Ecology & Management 66:512-528.

Type: Journal 

Geographic Area: North America

Keywords: carbon sequestration, land change science, 

social-ecological systems, social learning, sustainability, 

transformation

Abstract: Recent climatic trends and climate model projections indicate that climate change will modify rangeland 

ecosystem functions and the services and livelihoods that 

they provision. Recent history has demonstrated that climatic variability has a strong influence on both ecological 

and social components of rangeland systems and that these 

systems possess substantial capacity to adapt to climatic 

variability. Specific objectives of this synthesis are to: 1) 

evaluate options to mitigate greenhouse gas emissions and 

future climate change; 2) survey actions that individuals, 

enterprises, and social organizations can use to adapt to climate change; and 3) assess options for system transformation when adaptation is no longer sufficient to contend with 

climate change. Mitigation for carbon sequestration does not 

appear economically viable, given the small and highly variable carbon dioxide fluxes of rangeland ecosystems and the 

high transaction costs that would be incurred. In contrast, 

adaptation strategies are numerous and provide a means to 

manage risks associated with climate change. Adaptation 

strategies are diverse, including altered risk perception by 

individuals, greater flexibility of production enterprises, and 

modifications to social organizations that emphasize climatic 

variability, rather than consistency. Many adaptations represent ‘‘no regrets’’ actions because their implementation can 

be justified without emphasis on pending climate change. 

Adaptations specific to livestock production systems can include flexible herd management, alternative livestock breeds 

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 57

flow. For each, we review and propose ways that policy makers and managers can use evolutionary thinking to preserve 

threatened species, combat pest species, or reduce undesirable 

evolutionary changes. Such evolution-based management has 

potential to be a highly efficient and consistent way to create 

greater ecological resilience to widespread, rapid, and multifaceted environmental change.

Link: https://research.franklin.uga.edu/lankau/sites/research.

franklin.uga.edu.lankau/files/Lankau%20et%20al.%202011.

pdf

157. Leakey ADB, Ainsworth EA, Bernard SM, 

Markelz RJC, Ort DR, Placella SA, Rogers A, Smith 

MD, Sudderth EA, Weston DJ, Wullschleger SD, 

Yuan S. 2009. Gene expression profiling: opening the black 

box of plant ecosystem responses to global change. Global 

Change Biology 15:1201-1213.

Type: Journal 

Geographic Area: Global

Keywords: elevated CO2, genomic, microarray

Abstract: The use of genomic techniques to address ecological questions is emerging as the field of genomic ecology. 

Experimentation under environmentally realistic conditions 

to investigate the molecular response of plants to meaningful 

changes in growth conditions and ecological interactions is the 

defining feature of genomic ecology. Because the impact of 

global change factors on plant performance are mediated by 

direct effects at the molecular, biochemical, and physiological 

scales, gene expression analysis promises important advances 

in understanding factors that have previously been consigned 

to the “black box” of unknown mechanism. Various tools and 

approaches are available for assessing gene expression in 

model and nonmodel species as part of global change biology 

studies. Each approach has its own unique advantages and 

constraints. A first generation of genomic ecology studies in 

managed ecosystems and mesocosms have provided a testbed 

for the approach and have begun to reveal how the experimental design and data analysis of gene expression studies can be 

tailored for use in an ecological context.

Link: http://esd.lbl.gov/files/departments/ecology/publications/2009_GlobChangeBiol_Leakey_etal.pdf

158. Ledig FT, Kitzmiller JH. 1992. Genetic strategies 

for reforestation in the face of global climate change. Forest 

Ecology and Management 50:153-169.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: seed transfer, seed sources, local 

adaptation, forestry, assisted migration, translocation, uncertainty, review

155. Joyce LA, Blate GM, Littell J, McNulty SG, 

Millar CI, Moser S, Neilson RP, O’Halloran KA, 

Peterson DL. 2008. Adaptation options for climate-sensitive 

ecosystems and resources. Chapter 3. In: Julius SH, West JM, 

editors. Adaptation Options for Climate-Sensitive Ecosystems 

and Resources. Washington (DC): U.S. Environmental 

Protection Agency. p 1-66. 

Type: Government Document

Geographic Area: USA

Compilers’ Keywords: forest management, climate change, 

adaptive strategy, USFS, National Forest System

Summary: The National Forest System (NFS) is composed of 

155 national forests (NFs) and 20 national grasslands (NGs), 

which encompass a wide range of ecosystems, harbor much of 

the nation’s biodiversity, and provide myriad goods and services. The mission of the U.S. Forest Service (USFS), which 

manages the NFS, has broadened from water and timber to sustaining ecosystem health, diversity, and productivity to meet 

the needs of present and future generations. The evolution of 

this mission reflects changing societal values (e.g., increasing 

emphasis on recreation, aesthetics, and biodiversity conservation), a century of new laws, increasing involvement of the 

public and other agencies in NF management, and improved 

ecological understanding. Climate change will amplify the already difficult task of managing the NFS for multiple goals. 

This chapter offers potential adaptation approaches and management options that the USFS might adopt to help achieve its 

NF goals and objectives in the face of climate change.

Link: http://cfpub.epa.gov/ncea/cfm/recordisplay.

cfm?deid=180143

156. Lankau R, Jørgensen PS, Harris DJ, Sih A. 

2011. Incorporating evolutionary principles into environmental management and policy. Evolutionary Applications 

4:315-325.

Type: Journal 

Geographic Area: Global

Keywords: conservation biology, environmental management, evolution, gene flow, selection, variation

Abstract: As policymakers and managers work to mitigate 

the effects of rapid anthropogenic environmental changes, 

they need to consider organisms’ responses. In light of recent 

evidence that evolution can be quite rapid, this now includes 

evolutionary responses. Evolutionary principles have a long 

history in conservation biology, and the necessary next step for 

the field is to consider ways in which conservation policy makers and managers can proactively manipulate evolutionary processes to achieve their goals. In this review, we aim to illustrate 

the potential conservation benefits of an increased understanding of evolutionary history and prescriptive manipulation of 

three basic evolutionary factors: selection, variation, and gene 

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58 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

capacity limitations. These challenges are common to protected area agencies in developed countries and pervade those in 

developing countries, revealing that limited adaptive capacity 

represents a substantive barrier to biodiversity conservation 

and other protected area management objectives in an era of 

rapid climate change.

Link: http://www.ncbi.nlm.nih.gov/pubmed/21850505

160. Lempriere TC, Bernier PY, Carroll AL, 

Flannigan MD, Gilsenan RP, McKenney DW, Hogg 

EH, Pedlar J, Blain D. 2008. The importance of forest sector adaptation to climate change. Edmonton, Alberta, Canada: 

Natural Resources Canada, Canadian Forest Service, Northern 

Forestry Centre. NOR-X-416E. 78 p.

Type: Government Document

Geographic Area: Canada

Compilers’ Keywords: forest management, insects, greenhouse emissions, IPCC

Abstract: This report summarizes current knowledge about 

recent changes in the climate of Canada’s forests and projects further changes over this century based on scenarios of 

future global greenhouse gas emissions developed by the 

Intergovernmental Panel on Climate Change. Even with sustained reductions in global emissions the future climate is predicted to be quite different, meaning that adaptation will be 

essential. Impacts on the forest are already occurring and will 

be substantial in the future. The current upward trend in area 

burned annually is expected to continue. Forests will be prone 

to widespread stress induced by the changing climate, increasing the likelihood of pest outbreaks in the short to medium 

term. Recent outbreaks of several pests have exceeded in scope 

all previous known epidemics of these pests and are associated with the crossing of a climatic threshold. Invasion of the 

boreal forest by the mountain pine beetle, Dendroctonus ponderosae (Hopkins), appears likely, although the effect of this 

range expansion would likely be less severe than that observed 

recently in British Columbia, and outbreaks of the spruce budworm, Choristoneura fumiferana (Clemens), are predicted to 

be longer and more severe in the future. Future forest growth 

in response to climate change is expected to be variable; with 

growth reduction because of drought in parts of Canada’s western forests perhaps the most dramatic short- to medium-term 

outcome, though modestly increased growth in the east is predicted. Such impacts have implications for the cost and characteristics of timber supply and climate change will also affect 

forestry operations, recreation opportunities, biodiversity, and 

carbon storage. Planning based on past approaches will need to 

be reconsidered. Current objectives for sustainable forest management may not be attainable in the future, although there 

may be some new opportunities. Climate change may produce 

public safety risks, significant economic and social dislocation in forest-dependent communities including Aboriginal 

Abstract: If global warming materializes as projected, natural 

or artificial regeneration of forests with local seed sources will 

become increasingly difficult. However, global warming is far 

from a certainty and predictions of its magnitude and timing 

vary at least twofold, In the face of such uncertainty, reforestation strategies should emphasize conservation, diversification, 

and broader deployment of species, seed sources, and families. 

Planting programs may have to deploy non-local seed sources, 

imported from further south or from lower elevations, which 

necessitates a system for conserving native gene pools in seed 

banks or clone banks. Planting a diverse array of species or 

seed sources is a hedge against the uncertainty inherent in current projections of warming. Most tree improvement programs 

already stress genetic diversity and deployment of multiprogeny mixes, but may better prepare for climate change by 

testing selections in an even wider set of environments than is 

now the case.

Link: http://www.treesearch.fs.fed.us/pubs/24275

159. Lemieux CJ, Scott DJ. 2011. Changing climate, 

challenging choices: identifying and evaluating climate change 

adaptation options for protected areas management in Ontario, 

Canada. Environmental Management 48:675-690.

Type: Journal 

Geographic Area: Ontario, Canada

Keywords: climate change, parks, protected areas, adaptation, 

conservation, management, policy, Policy Delphi

Abstract: Climate change will pose increasingly significant 

challenges to managers of parks and other forms of protected 

areas around the world. Over the past two decades, numerous 

scientific publications have identified potential adaptations, 

but their suitability from legal, policy, financial, internal capacity, and other management perspectives has not been evaluated 

for any protected area agency or organization. In this study, a 

panel of protected area experts applied a Policy Delphi methodology to identify and evaluate climate change adaptation 

options across the primary management areas of a protected 

area agency in Canada. The panel identified and evaluated one 

hundred and sixty five (165) adaptation options for their perceived desirability and feasibility. While the results revealed 

a high level of agreement with respect to the desirability of 

adaptation options and a moderate level of capacity pertaining 

to policy formulation and management direction, a perception 

of low capacity for implementation in most other program 

areas was identified. A separate panel of senior park agency 

decision-makers used a multiple criterion decision-facilitation 

matrix to further evaluate the institutional feasibility of the 56 

most desirable adaptation options identified by the initial expert panel and to prioritize them for consideration in a climate 

change action plan. Critically, only two of the 56 adaptation 

options evaluated by senior decision-makers were deemed 

definitely implementable, due largely to fiscal and internal 

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 59

162. Maness TC. 2009. Forest management and climate change mitigation: good policy requires careful thought.

Journal of Forestry 107:119-124.

Type: Journal 

Geographic Area: USA

Keywords: climate change mitigation, forest management, 

forest policy

Abstract: Many people believe that forest management and 

the production of forest products can be important tools in a 

climate change mitigation strategy. This article critically examines how forest management can contribute to climate change 

mitigation in the global context, and it examines four major issues that must be considered before developing a strategy that 

would achieve real contributions. The article argues that protection of the carbon stock in the existing natural forest should 

be the central management objective related to carbon. Second 

should be the development of a widely accepted standard for 

calculating forestry offsets to drive future private investment 

on private forests. The article concludes with some recommendations for policy setting. 

163. Manter DK, Reeser PW, Stone JK. 2005.

A climate-based model for predicting geographic variation in Swiss needle cast severity in the Oregon Coast Range.

Phytopathology 95:1256-1265.

Type: Journal 

Geographic Area: Oregon, USA

Keywords: climate change, Douglas-fir, epidemiology, fungal 

pathogen, modeling

Abstract: Since the early 1990s, Swiss needle cast disease

caused by Phaeocryptopus gaeumannii has been increasing in 

Douglas-fir plantations in the Oregon Coast Range. Considerable variation in disease severity across the affected area often

has been noted. We investigated the influence of site microclimate on fungal colonization as a basis for this variation with a 

combination of seedling inoculation and field studies. Development of P. gaeumannii ascocarps on inoculated seedlings 

subjected to mist, irrigation, and shading treatments was 

followed for 10 months. Contrary to expectations, numbers

of ascocarps on foliage were negatively correlated with shade 

and mist and positively correlated with temperature. Numbers 

of ascocarps on foliage, site temperature, and leaf wetness 

were monitored over 5 years at nine field sites in the Oregon 

Coast Range. Factors most highly correlated with ascocarp

abundance were winter mean daily temperature and spring 

cumulative leaf wetness. Predictive models for disease severity

on the basis of these correlations were tested against disease 

and climate data measured at field sites during 2003–2004. A 

temperature-based disease prediction model was developed in 

combination with geographical information systems (GIS)-

linked climate databases to estimate disease levels across a 

communities, and impacts on the competitiveness of companies as well as on the actions and policies of all levels of government. These effects can be reduced through early identification and implementation of actions to reduce vulnerabilities or 

take advantage of new opportunities. The key needs associated 

with adaptation in the forest sector include awareness building 

and debate, improved knowledge and information, vulnerability assessments, planning frameworks and tools, and enhanced 

coordination and cooperation among governments and other 

forest sector participants. Meeting the challenge of adaptation 

will require sustained effort for many years.

Link: http://publications.gc.ca/collections/collection_2009/

nrcan/Fo133-1-416E.pdf

161. Lindenmayer DB, Steffen WL, Burbidge AA, 

Hughes L, Kitching RL, Musgrave W, Stafford 

Smith M, Werner PA. 2010. Conservation strategies 

in response to rapid climate change: Australia as a case study. 

Biological Conservation 143:1587-1593.

Type: Journal 

Geographic Area: Australia

Keywords: climate change, biodiversity conservation, 

Australia, climate responses, existing biodiversity stressors

Abstract: As in all parts of the globe, rapid climate change 

in Australia will have significant negative impacts on biodiversity. It also will interact with pre-existing stressors such as 

native vegetation clearing, altered natural disturbance regimes 

and invasive species—all of which already have major negative effects on biota in Australia. Strategies to reduce climate 

change impacts on Australian biodiversity include a mixture 

of mitigation and adaptation actions (sensu Millar et al. 2007) 

such as: (1) significantly reducing greenhouse gas emissions, 

(2) ensuring bio-diverse carbon capture, (3) better tackling preexisting stressors on biodiversity, (4) better preparing for the 

effects of major natural disturbances, (5) significantly improving off-reserve conservation efforts including fostering appropriate connectivity, and (6) enhancing the existing reserve 

system by making it more comprehensive, adequate and representative. The first strategy above demands a global response 

otherwise major mitigation attempts in Australia that are not 

paralleled elsewhere around the world will have little effect 

on climate change and, in turn, contribute little to enhanced 

biodiversity conservation. Strategies 2–6 demand multi-scaled 

responses, particularly at a regional level, given the major regional differences in direct climate change impacts and their interactions with pre-existing regional stressors. Well-developed 

multi-scaled conservation plans to implement these strategies 

currently do not exist, nor do appropriate institutional arrangements and capacities. Institutional reforms are urgently needed 

in Australia to develop the land management, monitoring and 

regional response capabilities required to conserve biodiversity on a continent already significantly modified.

CHAPTER 1 • CLIMATE CHANGE • STRATEGIES
60 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

approaches that promote reversible and incremental steps, and 

that favor ongoing learning and capacity to modify direction as 

situations change. We suggest that no single solution fits all future challenges, especially in the context of changing climates, 

and that the best strategy is to mix different approaches for 

different situations. Resources managers will be challenged to 

integrate adaptation strategies (actions that help ecosystems 

accommodate changes adaptively) and mitigation strategies 

(actions that enable ecosystems to reduce anthropogenic influences on global climate) into overall plans. Adaptive strategies 

include resistance options (forestall impacts and protect highly 

valued resources), resilience options (improve the capacity of 

ecosystems to return to desired conditions after disturbance), 

and response options (facilitate transition of ecosystems from 

current to new conditions). Mitigation strategies include options to sequester carbon and reduce overall greenhouse gas 

emissions. Priority-setting approaches (e.g., triage), appropriate for rapidly changing conditions and for situations where 

needs are greater than available capacity to respond, will become increasingly important in the future.

Link: http://www.treesearch.fs.fed.us/pubs/31774

166. Munang R, Rivington M, Takle ES, Mackey 

B, Thiaw I, Liu J. 2010. Climate information and capacity needs for ecosystem management under a changing climate. 

Procedia Environmental Sciences 1:206-227.

Type: Journal 

Geographic Area: Global

Keywords: ecosystem management, climate information, adaptation, ecosystem services, decision making

Abstract: The paper demonstrates the need to integrate across 

information types (i.e., weather, climate, socio-economic, policy and ecology) to better inform those involved in decisionmaking for ecosystem management. The provision of climate 

information and an understanding of ecosystem responses to 

climate change and variability urgently need to underpin any 

planning for the future. Integrating climatic information into 

risk assessment frameworks and adaptation planning is essential as it will enable better informed decision making in planning to ensure the adequate provision of ecosystem services 

(water, food, air quality, shelter etc.) and appropriate adaptation and mitigation strategies for the well being of both people 

and nature. A substantial mindset shift to fully recognize the 

fundamental role of ecosystems as life-supporting systems is 

urgently needed. The value given to ecosystems and the magnitude of effort to manage them has to be based on this mere 

fact and indeed, it should be an integral part of any climate 

change agreement.

167. Nagel L. 2013. Developing adaptive silvicultural strategies in the context of climate change. In: Forestry and Natural 

Resources Webinar Portal. USDA Forest Service. 91 p.

portion of the Oregon Coast Range. This model can be used 

for hypothesis testing and as a decision support tool for forest 

managers.

Link: http://www.ncbi.nlm.nih.gov/pubmed/18943355

164. Mawdsley JR, O’Malley R, Ojima DS. 2009. A 

review of climate-change adaptation strategies for wildlife management and biodiversity conservation. Conservation Biology 

23:1080-1089.

Type: Journal 

Geographic Area: Global

Keywords: adaptation, biodiversity, climate change, conservation, management, restoration, wildlife

Abstract: The scientific literature contains numerous descriptions of observed and potential effects of global climate change 

on species and ecosystems. In response to anticipated effects 

of climate change, conservation organizations and government 

agencies are developing “adaptation strategies” to facilitate the 

adjustment of human society and ecological systems to altered 

climate regimes. We reviewed the literature and climate-change 

adaptation plans that have been developed in United States, 

Canada, England, Mexico, and South Africa and found 16 general adaptation strategies that relate directly to the conservation 

of biological diversity. These strategies can be grouped into 

four broad categories: land and water protection and management; direct species management; monitoring and planning; 

and law and policy. Tools for implementing these strategies 

are similar or identical to those already in use by conservationists worldwide (land and water conservation, ecological restoration, agrienvironment schemes, species translocation, captive propagation, monitoring, natural resource planning, and 

legislation/regulation). Although our review indicates natural 

resource managers already have many tools that can be used 

to address climate-change effects, managers will likely need 

to apply these tools in novel and innovative ways to meet the 

unprecedented challenges posed by climate change.

Link: http://www.ncbi.nlm.nih.gov/pubmed/19549219

165. Millar CI, Stephenson NL, Stephens SL. 2007.

Climate change and forests of the future: managing in the face of 

uncertainty. Ecological Applications 17:2145-2151.

Type: Journal 

Geographic Area: USA

Keywords: carbon sequestration, climate change, desired 

conditions, ecosystem management, facilitated conservation, 

forest management, historical variability, resilience, resistance, 

wildfire

Abstract: We offer a conceptual framework for managing 

forested ecosystems under an assumption that future environments will be different from present but that we cannot be 

certain about the specifics of change. We encourage flexible 

CHAPTER 1 • CLIMATE CHANGE • STRATEGIES
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 61

Summary: The Strategy is the first joint effort of three levels 

of government (federal, state, and tribal) that have primary authority and responsibility for the living resources of the United 

States to identify what must be done to help these resources become more resilient, adapt to, and survive a warming climate. 

It is designed to inspire and enable natural resource managers, 

legislators, and other decision makers to take effective steps 

towards climate change adaptation over the next five to ten 

years. Federal, state, and tribal governments and conservation 

partners are encouraged to read the Strategy in its entirety to 

identify intersections between the document and their mission 

areas and activities. The Strategy is guided by nine principles. 

These principles include collaborating across all levels of government, working with non-government entities such as private landowners and other sectors like agriculture and energy, 

and engaging the public. It is also important to use the best 

available science—and to identify where science and management capabilities must be improved or enhanced. When adaptation steps are taken, it is crucial to carefully monitor actual 

outcomes in order to adjust future actions to make them more 

effective, an iterative process called adaptive management. We 

must also link efforts within the U.S. with efforts internationally to build resilience and adaptation for species that migrate 

and depend on areas beyond U.S. borders. Finally, given the 

size and urgency of the challenge, we must begin acting now.

Link: http://www.wildlifeadaptationstrategy.gov/strategy.php

169. Nitschke CR, Innes JL. 2008. Integrating climate change into forest management in South-Central British 

Columbia: an assessment of landscape vulnerability and development of a climate-smart framework. Forest Ecology and 

Management 256:313-327.

Type: Journal 

Geographic Area: Western Canada

Keywords: climate change, vulnerability, risk, forest management, triad zoning, modelling, fire, biodiversity, resilience, 

ecosystem, mountain pine beetle

Abstract: The achievement of sustainable forest management requires the incorporation of risk and uncertainty into 

long-term planning. Climatic change will have significant 

impacts on natural disturbances, species and ecosystems, 

particularly on landscapes influenced by forest management. 

Understanding where vulnerabilities lie is important in managing the risks associated directly or indirectly with climatic 

change. The vulnerability of landscapes to natural disturbances, the resilience of ecosystems and distribution of species are all important components that need to be considered 

when undertaking forest planning, but climatic change is rarely 

factored into such planning. In this study, the vulnerability of 

fire potential, fire regimes, ecosystems and species to climatic 

change was modelled for a 145,000 ha landscape in the southcentral interior of British Columbia, Canada. The results from 

Type: Presentation

Geographic Area: USA

Compilers’ Keywords: silviculture, adaptive strategy, forest 

management

Abstract: Climate change is a pervasive driver of ecosystem 

change and uncertainty regarding future conditions, and represents a significant threat to many critical ecosystem functions. 

The uncertain nature of climate change adds an additional 

dimension to developing sustainable management plans, including the development of silviculture prescriptions. Forest 

managers need conceptual tools to incorporate adaptation into 

management approaches. This process begins with reframing 

objectives and the notion of desired future conditions, which 

can be especially important where restoration treatments are 

a priority. Decision-making processes built on principles of 

adaptive management can help managers assess potential climate-related challenges to their goals and objectives, evaluate 

the feasibility of existing objectives in the context of climate 

change, diagnose the need for climate change adaptation efforts, prescribe appropriate treatments, and use monitoring efforts to evaluate treatment effectiveness and gauge the need to 

adjust management over time. A common emerging adaptation 

theme is to manage ecosystems for resistance and resilience, 

which can often be achieved by maintaining and enhancing 

complexity. A case study using northern hardwoods of the 

Great Lakes region, an extensive and economically important 

forest type that has undergone significant change and homogenization as a result of past land-use, will be discussed in the 

context of these tools. Additionally, an adaptive silviculture 

project will be highlighted that is building science-management partnerships to guide managers through the process of 

developing site-specific, objectives-driven climate change 

adaptation treatments. This project will simultaneously establish a framework for a long-term study of responses to several 

common climate change adaptation options in a variety of forest types across the United States.

Link: http://www.forestrywebinars.net/webinars/developingadaptive-silvicultural-strategies-in-the-context-of-climatechange

168. National Fish, Wildlife, and Plants Climate 

Adaption Partnership. 2012. National fish, wildlife and 

plants climate adaptation strategy. Washington (DC): Fish and 

Wildlife agencies, Council on Environmental Quality, Great 

Lakes Indian Fish and Wildlife Commission, National Oceanic 

and Atmospheric Administration, and U.S. Fish and Wildlife 

Service. 120 p.

Type: Government Document

Geographic Area: USA

Compilers’ Keywords: climate change, translocation, management planning

CHAPTER 1 • CLIMATE CHANGE • STRATEGIES
62 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

171. Obama B. 2013. The President’s climate action plan. 

Washington (DC): Executive Office of the President. 21 p.

Type: Government Document

Geographic Area: USA

Compilers’ Keywords: climate change, natural resources

Introduction: Climate change represents one of our greatest 

challenges of our time, but it is a challenge uniquely suited to 

America’s strengths. What follows is a blueprint for steady, 

responsible national and international action to slow the effects 

of climate change so we leave a cleaner, more stable environment for future generations. It highlights progress already set 

in motion by the Obama Administration to advance these goals 

and sets forth new steps to achieve them.

Link: http://www.whitehouse.gov/share/climate-action-plan

172. Ogden AE, Innes J. 2007. Incorporating climate 

change adaptation considerations into forest management 

planning in the boreal forest. International Forestry Review 

9:713-733.

Type: Journal 

Geographic Area: Canada

Keywords: climate change, adaptation, boreal forest, Montréal 

Process, forest management plans

Abstract: Climate change will pose increasing challenges to 

forest managers working to achieve sustainable forest management in the boreal forest. To date, discussions around when, 

where and how to consider adaptation in forest management 

plans for the boreal forest have been limited. As a starting 

point, specific objectives for climate change adaptation need to 

be articulated, which we consider to be synonymous with the 

criteria for conservation and sustainable management of boreal 

forests as defined by the Montréal Process. Secondly, because 

forest management plans are hierarchal—there are higher level 

strategic plans and lower level operational plans—it is important to distinguish at which planning level adaptation options 

are most appropriately considered. The purpose of this paper 

is to put forward a range of alternative adaptation options that 

forest managers working in the boreal zone could consider 

during the development of strategic and operational forest 

management plans in order to achieve sustainability as defined 

by the Montréal Process.

173. Oliver M. 2012. Adaptation: planning for climate 

change and its effects on Federal Lands. Portland (OR): USDA 

Forest Service, Pacific Northwest Research Station. Issue 148. 

6 p.

Type: Government Document

Geographic Area: USA

Compilers’ Keywords: adaptation strategy, forest management, USFS

these analyses were used to guide forest zoning, using the triad 

zoning framework, and for the development of a ‘‘climate 

smart’’ management framework. The use of climate-smart 

management is advocated as a decision-making framework 

for managing forested landscapes based on an understanding 

of landscape vulnerability to future climatic change. From this 

understanding, the maintenance of ecosystem health and vitality could be achieved.

Link: http://madsg.com/wp-content/uploads/2013/04/1-s2.0-

S037811270800368X-main.pdf

170. Northern Institute of Applied Climate 

Science [NIACS]. 2009. Climate change response framework. (URL accessed 25 November 2015)

Type: Web Page

Geographic Area: USA

Compilers’ Keywords: approach, forest management, forest 

trees, USFS

About: Resource managers face the immense challenge of 

integrating the inherent uncertainties of a changing climate 

into a wide variety of management decisions. These decisions occur along geographic scales, from several acres to 

millions of acres, and time scales, from immediate actions to 

long-term planning. Adaptation means taking action to enhance the ability of ecosystems to thrive in future conditions, 

but there is not a single answer for how to best adapt to climate change. Adaptation responses will vary by region based 

upon the magnitude of expected climate impacts, resilience 

of ecosystems, values and resources of local communities, 

and many other factors that demand unique responses to local conditions. The Climate Change Response Framework 

is a highly collaborative approach to helping land managers 

understand the potential effects of climate change on forest 

ecosystems and integrating climate change considerations 

into management. Since 2009, the Framework projects have 

worked to bridge the gap between scientific research on climate change impacts and on-the-ground management. The 

Framework stretches across the boundaries of partners to 

invite participation of forestlands owned and managed by 

private individuals, forest industry, tribes, state, local, and 

federal agencies. The Framework was initiated with a joint 

commitment of the U.S. Forest Service Northern Research 

Station and the Eastern Region to work closely together in 

addressing the challenges of climate change. The hallmark of 

the Framework, however, is the high level of cross-boundary 

cooperation among forest owners, considered essential to 

coping with an issue that spans borders, disciplines, and perspectives. There are currently three ecoregional Framework 

Projects that encompass 133 million acres in eight states, 

including 11 National Forests.

Link: http://www.climateframework.org/

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 63

altering native plant communities at a greater rate than previously anticipated and the effects on native plant communities could be extensive. To avoid the threat of habitats dominated by monocultures of invasive species, we may need to 

move and establish native plant materials to more northern 

latitudes if plant communities cannot adapt to climate change. 

Developing native plant materials and having native seed 

stored in long-term conservation storage and available on the 

market for restoration will provide federal agencies with some 

of the most important tools to help address threats to natural 

systems posed by destructive events such as wildfires, invasive 

species, and climate change.

Link: http://nativeseed.info/2010/presentations/Olwell.pdf

175. Peterson D, Millar CI, Joyce LA, Furniss MJ, 

Halofsky JE, Neilson RP, Morelli TL. 2011.Responding 

to climate change in national forests: a guidebook for developing adaptation options. Portland (OR): USDA Forest Service, 

Pacific Northwest Research Station. General Technical Report 

PNW-GTR-855. 109 p.

Type: Government Document

Geographic Area: USA

Keywords: adaptation, climate change, national forests, national parks, science-management partnership, vulnerability 

assessment

Abstract: This guidebook contains science-based principles, 

processes, and tools necessary to assist with developing adaptation options for national forest lands. The adaptation process 

is based on partnerships between local resource managers and 

scientists who work collaboratively to understand potential 

climate change effects, identify important resource issues, and 

develop management options that can capitalize on new opportunities and reduce deleterious effects. Because management objectives and sensitivity of resources to climate change 

differ among national forests, appropriate processes and tools 

for developing adaptation options may also differ. Regardless 

of specific processes and tools, the following steps are recommended: (1) become aware of basic climate change science 

and integrate that understanding with knowledge of local resource conditions and issues (review), (2) evaluate sensitivity 

of specific natural resources to climate change (rank), (3) develop and implement strategic and tactical options for adapting 

resources to climate change (resolve), and (4) monitor the effectiveness of adaptation options (observe) and adjust management as needed. Results of recent case studies on adaptation 

in national forests and national parks can facilitate integration 

of climate change in resource management and planning and 

make the adaptation process more efficient. Adaptation to climate change will be successful only if it can be fully implemented in established planning processes and other operational 

aspects of national forest management.

Link: http://www.treesearch.fs.fed.us/pubs/39884

Summary: National forest managers are charged with tackling the effects of climate change on the natural resources 

under their care. The Forest Service National Roadmap for 

Responding to Climate Change and the Climate Change 

Performance Scorecard require managers to make significant 

progress in addressing climate change by 2015. To help land 

managers meet this challenge, Forest Service scientists conducted three case studies on national forests and adjacent national parks and documented a wide range of scientific issues 

and solutions. They summarized the scientific foundation for 

climate change adaptation and made the information accessible to land managers by creating a climate change adaptation 

guidebook and web portal. Case study teams discovered that 

collaboration among scientists and land managers is crucial 

to adaptation planning, as are management plans targeted to 

the particular ecosystem conditions and management priorities of each region. Many current management practices are 

consistent with climate change adaptation goals. Because 

timely implementation is critical, strategies are in development 

at the national level to speed the implementation of sciencebased climate change adaptation processes in national forests 

throughout the country.

Link: http://www.treesearch.fs.fed.us/pubs/42317

174. Olwell P. 2010. National native plant materials development program: ensuring options in a changing climate. 

National Native Seed Conference: Native Plant Materials 

Development, Production and Use in Habitat Restoration. 

Corvallis (OR): Institute for Applied Ecology. 40 p.

Type: Presentation

Geographic Area: USA

Compilers’ Keywords: seeds of success, seed storage, migration, BLM

Abstract: Following record breaking wildfire seasons of 1999 

and 2000, Congress directed the Bureau of Land Management 

(BLM) to develop and implement a program for site appropriate native plant materials. The NPMDP is coordinating organizations around the country to collect, curate and conserve 

plant diversity before it is lost. With the assistance of more 

than 500 partners, BLM is leading the interagency Native 

Plant Materials Development Program (NPMDP) to ensure 

the quality and quantity of genetically appropriate native plant 

materials are available commercially for restoring native plant 

communities across the American landscape. Developing 

a crop from native wild species begins with seed collection. 

Seeds of Success (SOS) is the native seed collection phase of 

the NPMDP. SOS makes the primary collection of common 

native species that are needed for restoration after fire, weed 

eradication, energy development, or for use in climate adaptation strategies. Almost 100 teams are collecting nation-wide to 

add to the more than 9,500 collections which collectively form 

the Seeds of Success National Collection. Climate change is 

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64 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Abstract: Conservation planning is the process of locating, configuring, implementing and maintaining areas that 

are managed to promote the persistence of biodiversity and 

other natural values. Conservation planning is inherently 

spatial. The science behind it has solved important spatial 

problems and increasingly influenced practice. To be effective, however, conservation planning must deal better with 

two types of change. First, biodiversity is not static in time 

or space but generated and maintained by natural processes. 

Second, humans are altering the planet in diverse ways at 

ever faster rates.

Link: http://www.ncbi.nlm.nih.gov/pubmed/17981360

178. Rowland EL, Davison JE, Graumlich LJ. 

2011. Approaches to evaluating climate change impacts on 

species: a guide to initiating the adaptation planning process.

Environmental Management 47:322-337.

Type: Journal 

Geographic Area: Global

Keywords: climate change, vulnerability assessments, management, species, adaptation planning 

Abstract: Assessing the impact of climate change on species and associated management objectives is a critical initial step for engaging in the adaptation planning process. 

Multiple approaches are available. While all possess limitations to their application associated with the uncertainties 

inherent in the data and models that inform their results, 

conducting and incorporating impact assessments into the 

adaptation planning process at least provides some basis for 

making resource management decisions that are becoming 

inevitable in the face of rapidly changing climate. Here we 

provide a non-exhaustive review of long-standing (e.g., species distribution models) and newly developed (e.g., vulnerability indices) methods used to anticipate the response to 

climate change of individual species as a guide for managers 

grappling with how to begin the climate change adaptation 

process. We address the limitations (e.g., uncertainties in 

climate change projections) associated with these methods, 

and other considerations for matching appropriate assessment approaches with the management questions and goals. 

Thorough consideration of the objectives, scope, scale, time 

frame and available resources for a climate impact assessment allows for informed method selection. With many data 

sets and tools available on-line, the capacity to undertake 

and/or benefit from existing species impact assessments is 

accessible to those engaged in resource management. With 

some understanding of potential impacts, even if limited, adaptation planning begins to move toward the development of 

management strategies and targeted actions that may help to 

sustain functioning ecosystems and their associated services 

into the future.

Link: http://www.ncbi.nlm.nih.gov/pubmed/21259061

176. Pojar J. 2010. A new climate for conservation: nature, carbon and climate change in British Columbia. British 

Columbia, Canada: Working Group on Biodiversity, Forests 

and Climate. 100 p.

Type: Report

Geographic Area: Western Canada

Compilers’ Keywords: strategy, carbon management

Introduction: A New Climate for Conservation: Nature, 

Carbon and Climate Change in British Columbia explores the 

role of nature conservation in a climate action strategy for ecological adaptation (Part 1) and ecological mitigation (Part 2), 

with the key recommendation to develop a comprehensive and 

integrated Nature Conservation and Climate Action Strategy 

for the Province of British Columbia (Part 3): Part 1 presents 

available science on current climate-change projections, and 

present and future impacts of climate change to ecosystems, 

species, genotypes, and the processes Linking them. The review focuses primarily on forested systems, and also addresses 

non-forest and aquatic systems. Ecosystem resilience and adaptation options, in relation to climate change, are outlined. 

Current thinking in conservation science is then summarised 

in light of external pressures. B.C.’s existing conservation 

planning and forestry management are reviewed in terms of 

their ability to respond to the challenges of climate change. 

Part 2 summarises literature on natural capital, ecosystem 

services and the role of ecosystems in climate change mitigation. Variations in carbon sequestration and storage in different 

ecosystems are discussed and research gaps in forest carbon 

dynamics are identified. Current opportunities for an offset 

market through carbon activities such as avoided degradation, ecological restoration and improved forest management 

are also explored, in light of recent pilot projects in B.C. Part 

3 integrates the findings from Part 1 and Part 2 in a central 

recommendation—to develop a comprehensive and integrated 

provincial Nature Conservation and Climate Action Strategy. 

To be efficient, this strategy must combine nature conservation 

and carbon/climate management planning. To be effective, it 

must embrace the fundamental role of conserving natural ecosystems for adaptation and mitigation of climate change and 

for nature’s many other ecosystem services, which underpin 

sustainable options for current and future generations.

Link: http://www.borealbirds.org/resources/report-pojarbcconservation.pdf

177. Pressey RL, Cabeza M, Watts ME, Cowling 

RM, Wilson KA. 2007. Conservation planning in a changing world. Trends in Ecology and Evolution 22:583-592.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: animals, biodiversity, humans, social 

change

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 65

case studies that highlight how biodiversity conservation can 

be used in planning. We conclude with general guidance on 

choosing appropriate climate adaptation approaches to amend 

for conservation planning.

Link: http://www.bioone.org/doi/abs/10.3375/043.035.0120

181. Schwartz MW. 2012. Using niche models with 

climate projections to inform conservation management 

decisions. Biological Conservation 155:149-156.

Type: Journal 

Geographic Area: Global

Keywords: species distribution model, niche model, climate 

change, resource management, adaptive management, decision theory, extinction, natural resource management

Abstract: Conservation science strives to inform management decisions. Applying niche models in concert with future climate projections to project species vulnerability to 

extinction, range size loss, or distribution shifts has emerged 

as a potentially useful tool for informing resource management decisions. Making climate change niche modeling 

useful to conservation decisions requires centering studies 

on the types of decisions that are made regarding the focal 

taxa of a niche model study. Recent recommendations for 

climate adaptation strategies suggest four types of decision 

makers: policy, habitat protection, habitat management, species management. Targeting research to questions relevant 

for management decisions will increase utility of a niche 

model study. Constraints to the accuracy and precision of 

niche models to project potential future distributions are 

well recognized. How to incorporate these uncertainties into 

management decision-making remains a challenge. Refining 

estimates and making sound management recommendations 

is critical because species that are generally modeled to be 

the most vulnerable to climate change (i.e., narrow endemics), are also the most vulnerable to bad decisions based on 

uncertain models. I review uncertainties of niche models 

to assert that there is an inherent bias for models to overestimate climate-driven vulnerability to extirpation. Explicit 

recognition of this bias leads to a decision framework that 

accommodates unbalanced uncertainty. Namely, niche models may be more useful for identifying conservation opportunities identifying newly available habitats under changing 

climate than they are for asserting where current habitat will 

no longer exist under future climate states.

182. Seabrook L, McAlpine CA, Bowen ME. 2011.

Restore, repair or reinvent: options for sustainable landscapes in a changing climate. Landscape and Urban Planning 

100:407-410.

Type: Journal 

Geographic Area: Global

179. Schmidtling RC. 1994. Use of provenance tests to 

predict response to climatic change: Loblolly pine and Norway 

spruce. Tree Physiology 14:805-817.

Type: Journal 

Geographic Area: Southeastern USA

Keywords: genetic response, growth, height growth, seed 

source, temperature, tree response 

Summary: Provenance tests are often used to determine genetic responses of seed sources to transfer to different climates. 

This study was undertaken to determine whether provenance 

tests can be used to predict tree response to rapid climate 

changes in situ. Data from provenance tests of loblolly pines 

(Pinus taeda L.), Norway spruce (Picea abies L. Karst) and 

other southern pines (subsect. AUSTRALES Loud.) were interpreted using regression models to relate growth to temperature variables. Results of different plantings were combined 

by expressing growth as a percent deviation from the “local” 

source, and expressing temperature at the source as a deviation 

from that of the planting site. The results of the loblolly pine 

and Norway spruce models predicted a loss of about 5 to 10% 

in height growth below that expected for a genetically adapted 

seed source, if the average yearly temperature increases by 

4 °C.

Link: http://treephys.oxfordjournals.org/content/14/7-8-9/805.full.pdf

180. Schmitz OJ, Lawler JJ, Beier P, Groves C, 

Knight G, Boyce DA, Bulluck J, Johnston KM, Klein 

ML, Muller K, Pierce DJ, Singleton WR, Strittholt 

JR, Theobald DM, Trombulak SC, Trainor A. 2015. 

Conserving biodiversity: practical guidance about climate 

change adaptation approaches in support of land-use planning.

Natural Areas Journal 35:190-203.

Type: Journal

Geographic Area: USA

Keywords: biodiversity, climate adaptation, conservation 

planning, land use planning

Abstract: As species’ geographic ranges and ecosystem 

functions are altered in response to climate change, there is 

a need to integrate biodiversity conservation approaches that 

promote natural adaptation into land use planning. Successful 

conservation will need to embrace multiple climate adaptation 

approaches, but to date they have not been conveyed in an 

integrated way to help support immediate conservation planning and action in the face of inherent spatial uncertainty about 

future conditions. Instead, these multiple approaches are often 

conveyed as competing or contradictory alternatives, when in 

fact, they are complementary. We present a framework that 

synthesizes six promising spatially explicit adaptation approaches for conserving biodiversity. We provide guidance 

on implementing these adaptation approaches and include 

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66 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

decisions, and putting the emerging field of restoration genetics into practice.

Link: http://riverpartners.org/documents/press_releases/

PressRelease_20090831.pdf

184. Sgrò CM, Lowe AJ, Hoffmann AA. 2011.

Building evolutionary resilience for conserving biodiversity under climate change. Evolutionary Applications 4:326-337.

Type: Journal 

Geographic Area: Global

Keywords: adaptive potential, biodiversity, climate change, 

conservation, evolution, evolutionary resilience, genetic 

diversity

Abstract: Evolution occurs rapidly and is an ongoing process 

in our environments. Evolutionary principles need to be built 

into conservation efforts, particularly given the stressful conditions organisms are increasingly likely to experience because 

of climate change and ongoing habitat fragmentation. The concept of evolutionary resilience is a way of emphasizing evolutionary processes in conservation and landscape planning. 

From an evolutionary perspective, landscapes need to allow 

in situ selection and capture high levels of genetic variation 

essential for responding to the direct and indirect effects of 

climate change. We summarize ideas that need to be considered in planning for evolutionary resilience and suggest how 

they might be incorporated into policy and management to ensure that resilience is maintained in the face of environmental 

degradation.

Link: http://www.trend.org.au/sites/default/files/Sgro_Lowe_

Hoffman_2011.pdf

185. Shoo LP, Hoffmann AA, Garnett ST, Pressey 

RL, Williams YM, Taylor M, Falconi L, Yates CJ, 

Scott JK, Alagador D, Williams SE. 2013. Making 

decisions to conserve species under climate change. Climatic 

Change 119:239-246.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: biodiversity, decision framework, 

assisted colonization

Abstract: Severe impacts on biodiversity are predicted to arise 

from climate change. These impacts may not be adequately addressed by conventional approaches to conservation. As a result, additional management actions are now being considered. 

However, there is currently limited guidance to help decision 

makers choose which set of actions (and in what order) is most 

appropriate for species that are considered to be vulnerable. 

Here, we provide a decision framework for the full complement of actions aimed at conserving species under climate 

change from ongoing conservation in existing refugia through 

various forms of mobility enhancement to ex situ conservation 

Keywords: restoration, novel ecosystems, landscape change, 

ecosystem function

Abstract: Globally, land use and land cover changes to provide resources for human society have led to loss of biodiversity and declines in ecosystem services. Climate change will 

compound these impacts. Ecological restoration can reverse 

environmental degradation and is starting to restore ecological 

services, as well as help mitigate climate change. Although this 

may be an anathema to some, we must systematically assess 

and proactively redesign and manage the landscapes we inhabit so they can continue to provide ecosystem services essential 

for all species, including humans. We define three restoration 

pathways based largely on existing land use and the degree 

of modification: restoration for areas of natural ecosystems; 

repair for production landscapes; and reinvention for urban 

areas. Challenges and research priorities include understanding: the effects of mean climate change and climate extremes 

on species’ distribution and ecosystem composition; how restoration can be used for carbon sequestration; the effects of 

proximate and endogenous drivers on landscape change; how 

to better bio-design landscapes for multiple functions; integration of different scales of restoration planning and design; 

and the establishment of long-term monitoring and adaptive 

management.

183. Seavy NE, Gardali T, Golet GH, Griggs FT, 

Howell CA, Kelsey R, Small SL, Viers JH, Weigand 

JF. 2009. Why climate change makes riparian restoration 

more important than ever: recommendations for practice and 

research. Ecological Restoration 27:330-338.

Type: Journal 

Geographic Area: California, USA

Keywords: California, climate change, genetics, hydrology, 

restoration, riparian

Abstract: Over the next century, climate change will dramatically alter natural resource management. Specifically, historical reference conditions may no longer serve as benchmarks 

for restoration, which may foster a “why bother?” attitude toward ecological restoration. We review the potential role for riparian restoration to prepare ecological systems for the threats 

posed by climate change. Riparian ecosystems are naturally 

resilient, provide linear habitat connectivity, link aquatic and 

terrestrial ecosystems, and create thermal refugia for wildlife: 

all characteristics that can contribute to ecological adaptation 

to climate change. Because riparian systems and the projected 

impacts of climate change are highly variable geographically, 

there is a pressing need to develop a place-based understanding 

of climate change threats to riparian ecosystems. Restoration 

practitioners should consider how they can modify practices 

to enhance the resilience of riparian ecosystems to climate 

change. Such modifications may include accelerating the restoration of private lands, participating in water management 

CHAPTER 1 • CLIMATE CHANGE • STRATEGIES
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 67

management is well documented, there remain critical gaps in 

the portfolio of evidence. One of these concerns the impacts 

of germplasm collection, conservation, characterization and 

evaluation and the availability of evidence to determine impacts. Compared to literature about the impacts of CGIAR 

crop breeding programs, the body of published literature about 

the costs and benefits of gene bank collections and the services 

they provide is both small and narrow in the value components 

measured. The more theoretical work in this literature has 

also been of limited relevance to the problems and constraints 

faced by gene bank managers and system-wide bank system. 

Some of the gray literature produced by experts associated 

closely with the CGIAR gene banks has been more useful 

from an operational standpoint, but cannot yet be generalized 

across crops and collections because of important differences 

among the cost and benefits of conserving various crops due to 

their characteristics, bank location, and divergent operational 

approaches.

Link: http://impact.cgiar.org/sites/default/files/images/SmaleHansen2011.pdf

188. Solomon AM, Birdsey R, Joyce L, Hayes J. 

2009. Forest Service global change research strategy, 2009-

2010. Washington (DC): USDA Forest Service, Research and 

Development. FS-917a. 20 p.

Type: Government Document

Geographic Area: USA

Compilers’ Keywords: climate change, adaptive strategy, 

framework, research and development, forest management, 

USFS

Summary: In keeping with the research goals of the 

U.S. Climate Change Science Program, the Research and 

Development agenda of the Forest Service, USDA helps define 

climate change policy and develop best management practices 

for forests (both rural and urban) and grasslands. These actions 

are taken to sustain ecosystem health, adjust management for 

ecosystem services (“adaptation”), and increase carbon sequestration (“mitigation”), all under changing climate conditions. The fundamental research focus of the Forest Service 

Global Change Research Strategy is to increase understanding 

of forest, woodland, and grassland ecosystems so that they can 

be managed in a way that sustains and provides ecosystem services for future generations.

Link: http://www.treesearch.fs.fed.us/pubs/41132

189. Spittlehouse DL. 2005. Integrating climate change 

adaptation into forest management. The Forestry Chronicle 

81:691-695.

Type: Journal 

Geographic Area: Canada

outside the natural environment. We explicitly recognize that 

allocation of conservation resources toward particular actions 

may be governed by factors such as the likelihood of success, 

cost and likely co-benefits to non-target species in addition to 

perceived vulnerability of individual species. As such, we use 

expert judgment of probable tradeoffs in resource allocation 

to inform the sequential evaluation of proposed management 

interventions.

186. Sjølie HK, Latta GS, Solberg B. 2013. Potentials 

and costs of climate change mitigation in the Norwegian forest 

sector: does choice of policy matter? Canadian Journal of Forest 

Research 43:589-598.

Type: Journal 

Geographic Area: Norway, Europe

Compilers’ Keywords: forest management, greenhouse 

gases, emissions

Abstract: Forests are important contributors to the global 

carbon cycle and mitigate climate change through carbon sequestration and the supply of wood that substitutes for fossil 

fuels and greenhouse gas (GHG)-intensive building materials. 

However, current climate policies only partially credit forest carbon sequestration and bioenergy policies are handled 

independently of forestry. Using Norway as a case study, we 

analyze two sets of simulated carbon tax/subsidy policies, one 

crediting forest carbon sequestration while maintaining predetermined harvest levels and utilization of wood, and another 

targeting GHG fluxes in the entire forest industrial sector allowing harvest levels and wood markets to change in response 

to the policy. Results indicate that GHG emission reduction 

potentials differ substantially between the two policies, being 

several times higher for the latter than the former policy at a 

given carbon price. This suggests that (i) previous research efforts in Europe have not captured the full mitigation potential 

as they have not included adaptations in the harvest level and 

the wood market and (ii) climate policies should target GHG 

fluxes in the entire sector to utilize its potential contribution for 

mitigating climate change.

187. Smale M, Hanson J. 2010. Assessing the impact 

of CGIAR investments in germplasm collection, conservation, 

characterization and evaluation: the scope of the literature.

Consultative Group on International Agricultural Research. 

37 p.

Type: Report

Geographic Area: Global

Compilers’ Keywords: germplasm, literature review, 

Consultative Group on International Agricultural Research

Introduction: The terms of reference for the study to which 

this paper contributes state that while the impact of CGIAR 

research in crop germplasm improvement and integrated pest 

CHAPTER 1 • CLIMATE CHANGE • STRATEGIES
68 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

191. St. Clair JB, Howe GT. 2009. Genetic options for 

adapting forests to climate change. Western Forester 54:9-11.

Type: Newsletter

Geographic Area: North America

Compilers’ Keywords: restoration, forest genetics, local 

seed sources, management options

Introduction: Concerns over impacts of climate change 

on forest productivity and health, and questions about appropriate management responses, led to the formation of 

the Taskforce on Adapting Forests to Climate Change. An 

accompanying article discussed silvicultural approaches 

to deal with climate change. This article discusses genetic 

options including management actions that could be taken 

to influence the natural or human selection of genotypes, 

movement of genotypes across the landscape and conservation of genetic diversity. 

Link: http://www.treesearch.fs.fed.us/pubs/34501

192. St. Clair JB, Howe GT. 2011. Strategies for conserving forest genetic resources in the face of climate change. 

Turkish Journal of Botany 35:403-409.

Type: Journal 

Geographic Area: North America

Keywords: climate change, genetic conservation, adaptation

Abstract: Conservation of genetic diversity is important for 

continued evolution of populations to new environments, as 

well as continued availability of traits of interest in genetic 

improvement programs. Rapidly changing climates present 

new threats to the conservation of forest genetic resources. 

We can no longer assume that in situ reserves will continue to preserve existing genetic diversity. Management 

of reserves should become more active. In some reserves, 

existing genetic diversity should be preserved by creating 

stands that are more resistant to threats using silvicultural 

treatments such as thinning and prescribed burning. In other 

reserves, natural selection and adaptation to changed environments should be promoted by increasing within population genetic diversity and promoting gene flow. This may 

be done by locating reserves in areas of high environmental 

heterogeneity, minimizing fragmentation, and using assisted 

colonization to increase genetic diversity by establishing 

populations adapted to future climates within or adjacent to 

reserves. Threats to native stands from climate change and 

other interacting threats should bring a renewed importance 

to ex situ collections, particularly for rare and disjunct populations and those at the warmer and drier edges of a species 

range. Assisted colonization to move threatened populations 

to new environments must be considered as an additional 

conservation measure.

Link: http://www.treesearch.fs.fed.us/pubs/39973

Keywords: climate change, impacts, adaptation, vulnerability, forests, ecosystems, risk management

Abstract: Future climate change will affect society’s ability to use forest resources. We take account of climate in 

forest management and this will help us adapt to the effects 

of climate change on forests. However, society will have to 

adjust to how forests adapt by changing expectations for the 

use of forest resources because management can only influence the timing and direction of forest adaptation at selected 

locations. There will be benefits as well as loses and an important component of adaptation will be balancing values. 

Adaptation options to respond to impacts on the timber supply in Canada for the next 50 to 100 years are limited mainly 

to forest protection and wood utilisation because these forests are already in the ground. Adaptation through reforestation will focus on commercial tree species. It is important 

to start developing adaptation strategies now. These include 

assessing forest vulnerability to climate change, revising 

expectations of forest use, determining research and educational needs, development of forest policies to facilitate 

adaptation, and determining when to implement responses. 

Government agencies should take the lead in creating an environment to foster adaptation in forestry and in developing 

the necessary information required to respond.

190. Spittlehouse DL, Stewart RB. 2003. Adaptation 

to climate change in forest management. BC Journal of 

Ecosystems and Management 4:1-11.

Type: Journal 

Geographic Area: North America

Keywords: climate change, sustainable forest management, 

adaptation

Abstract: Adaptation in forestry is sustainable forest management that includes a climate change focus. Climate 

change over the next 100 years is expected to have significant impacts on forest ecosystems. The forestry community 

needs to evaluate the long-term effects of climate change on 

forests and determine what the community might do now 

and in the future to respond to this threat. Management can 

influence the timing and direction of forest adaptation at selected locations, but in many situations society will have to 

adjust to however forests adapt. Adapting to climate change 

in the face of the uncertain timing of impacts means we must 

have a suite of readily available options. A high priority will 

be coping with and adapting to forest disturbance while 

maintaining the genetic diversity and resilience of forest 

ecosystems. A framework for facilitating adaptation in forestry is discussed and a review of adaptive actions presented.

Link: http://citeseerx.ist.psu.edu/viewdoc/download?doi=10.1

.1.170.1072&rep=rep1&type=pdf

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 69

Some questions pertained to the indicator’s relationships with 

other indicators in the set, some to the expected influences of 

climate change on the entity represented by the indicator, and 

finally some that would help us understand the indicator’s ongoing relevance to SFM under a changing climate. The research 

team consisted of experienced forestry professionals and researchers who applied their collective professional judgment, 

as informed by a thorough canvassing of relevant literature, in 

answering the questions and developing recommendations for 

each of the indicators. The preliminary findings, and indeed the 

entire report, were peer-reviewed by experts from across Canada. 

Forty-six indicators were examined using the evaluation protocol. The findings are summarized in the main report and detailed 

in a companion report. The evaluated indicators were assigned 

to one of three general outcomes. Twelve indicators were considered to be entirely independent of climate change, meaning 

that climate change is not expected to affect the phenomena 

represented by these indicators. The utility and robustness of 

all the remaining 34 indicators was considered to be influenced 

by a changing climate. For 23 of these, the team recommends 

no change to the indicator (unmodified category), and for 11 of 

them, changes are recommended. Initially it was thought that a 

potential outcome of the evaluation could be outright abandonment of an indicator in the event that the team found its ongoing 

utility to be seriously eroded by climate change. However, none 

of the indicators was found in this situation. Finally, he study 

identified six new indicators that could help provide a climatechange lens for monitoring and managing forests sustainably 

in Canada. These are: a) Connectivity of protected areas; b) 

Proportion of tenured forest area with seed transfer guidelines 

that account for climate change; c) Average, minimum, and 

maximum temperature d) Area of Crown forest with assisted 

migration initiatives; e) Rate and form of precipitation; and f) 

Carbon emissions avoided through product substitution The 

study concludes with a set of recommendations that should help 

improve the overall utility of the C&I-SFM, especially in the 

context of a changing climate. These recommendations address: 

(a) moving from predominantly retrospective analysis using 

C&I-SFM to a balance of retrospective and prospective analysis; (b) Linking C&I-SFM much more strongly and directly into 

forest management and policy processes; (c) undertaking analytical work using a framework of complex adaptive systems; 

(d) making explicit consideration of climate change in all forest 

management and policy decisions; and (e) sector-wide collaboration in ongoing improvement to and application of the C&ISFM. The report concludes by reminding readers that C&I-SFM 

are a necessary element of the SFM enterprise. Progress is nigh 

impossible without using them. Confidence in such progress is 

indeed impossible without them. It is clear that climate change 

will affect the entire forest sector, sometimes in insidious ways, 

sometimes in abrupt and obvious ways, and sometimes even in 

helpful ways. Considering the complex manner in which climate 

change will interact with other human influences on forests and 

the sector, incisive cumulative effects assessment will become 

193. St. Clair JB, Howe GT, Erickson VJ. 2010.

Responding to climate change: genetic options. USDA Forest 

Service Genetic Resource Management Climate Change 

Workshop. Corvallis, OR. 70 p.

Type: Presentation

Geographic Area: Western USA

Compilers’ Keywords: Douglas-fir, Pseudotsuga menziesii, 

provenance test, seed transfer guidelines, seed transfer zones

Introduction: When considering ecosystem and management responses to climate change, it is important to consider 

genetics of adaptation and genetic variation in adaptive traits. 

There are three considerations: 1. Plants are genetically adapted to their local climates. The climatic tolerances of populations are considerably lower than the tolerances of the species 

as a whole. Populations, not species, are the important biological unit of interest. 2. Evolutionary adaptation will determine 

what happens to plant populations given climate change. 3. 

Management of genetic variation may positively influence 

how plants respond and adapt to climate change.

Link: http://www.fs.fed.us/wwetac/projects/erickson.html

194. Steenberg JWN, Duinker PN, Van Damme 

L, Zielke K. 2011. Indicators of sustainable forest management in a changing climate. Ottawa, Ontario, Canada: Canadian 

Council of Forest Ministers. 51 p.

Type: Government Document

Geographic Area: Canada

Compilers’ Keywords: climate change, Ontario, 

sustainability

Summary: The theme of sustainability is now woven throughout Canadian forest management and policy. Indeed, Canada 

was an early adopter of sustainability in forestry, largely through 

a series of initiatives led by the Canadian Council of Forest 

Ministers’ (CCFM). One such initiative was development and 

application of a national suite of criteria and indicators of sustainable forest management (C&I-SFM). With appropriate data 

associated with each of the chosen indicators, the C&I-SFM 

serve up a comprehensive picture of forest and forest-sector 

progress on the road called sustainable development. For almost 

two decades, C&I-SFM have been applied at a wide range of 

levels, from international to national to provincial to local. The 

national set for Canada has been revised once (2003), and a second revision is likely imminent. In the meantime, the CCFM, 

in its 2008 vision for Canada’s forests, stated that all initiatives 

related to SFM must consider the implications of a changing 

climate. In the context of a revision to the C&I-SFM, the question becomes this: what are the implications of a changing climate for the ongoing robustness and utility of Canada’s national 

C&I-SFM? This report attempts to address that question. The 

approach consisted of devising a systematic set of questions, 

in other words, an evaluation protocol, to put to each indicator. 

CHAPTER 1 • CLIMATE CHANGE • STRATEGIES
70 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

to change (short-term adaptation), Promote resilience to 

change (medium-term adaptation), Enable ecosystems to respond to change (long-term adaptation) In grazing management, most of the adaptations that have been discussed are 

based on short-term resistance to drought or other extreme 

events. These include reducing cattle herds, finding alternative grazing, increased feeding, and addressing stockwater 

shortages. Medium-term adaptations aimed at increasing the 

resilience of grazing operations include moderate to conservative stocking rates, maintaining litter cover, more flexible 

herd structure, and improving water supply systems. Longterm response options in grazing management include grassland monitoring, revision of range management standards, 

changes in grazing and land use strategies, and increases in 

management flexibility. In biodiversity conservation, most 

of the adaptations that have been discussed are medium- to 

long-term resilience and response strategies. These include 

incorporating climate change into conservation plans, increasing protected areas, mitigating other threats to biodiversity such as exotic invasion, and improving information 

on climate change. The strategies most focused on longterm response aim at facilitating the northward movement 

of species to adjust to the warmer climate. There are two 

broad approaches, with advantages and disadvantages in 

different situations: increasing landscape connectivity, and 

assisted migration of selected species. Existing government 

programs that directly or indirectly address climate change 

in grassland management are reviewed. It is recommended 

that government agencies use the list of adaptation options 

provided here to review their current policies and programs, 

with the aim of identifying those that could be modified or 

expanded to better address climate change. Options that are 

not addressed by any current policies should be the focus of 

new policy development.

Link: http://www.cakex.org/sites/default/files/

documents/12855-1E12%20Adaptation%20to%20ClimateChange%20in%20Grassland%20Management.pdf

197. Toth FL, Mwandosya M. 2001. Decision-making 

frameworks. Chapter 10. In: Metz B, Davidson O, Swart R, 

Pan J, editors. Climate Change 2001: Mitigation. Cambridge 

(MS): Cambridge University Press. p 601-688. 

Type: Book Section

Geographic Area: Global

Compilers’ Keywords: climate change, IPCC, mitigation, 

management, review

Summary: This chapter is a review of decision-making framework as applied to climate change mitigation. By taking into 

account the broader perspective of sustainable development 

the portfolio of mitigation policies is enhanced. A central issue in linking development and climate concerns is technological transfer that could help less-developed countries speed-up 

increasingly important. Rigorous application of C&I-SFM will 

help develop the insight needed to assess the real prospects for 

SFM in Canada under a changing climate.

Link: http://www.for.gov.bc.ca/ftp/hfp/external/!publish/web/

ffesc/reports/CI-SFM-ClimateChange_FinalReport_111017.

pdf

195. Strange N, Thorsen BJ, Bladt J, Wilson KA, 

Rahbek C. 2011. Conservation policies and planning under 

climate change. Biological Conservation 144:2968-2977.

Type: Journal 

Geographic Area: Global

Keywords: climate change, dynamic, policy, reserve selection, uncertainty

Abstract: Biodiversity conservation policies focus on securing the survival of species and habitats according to their current distribution. This basic premise may be inappropriate for 

halting biodiversity decline under the dynamic changes caused 

by climate change. This study explores a dynamic spatial conservation prioritization problem where climate change gradually changes the future habitat suitability of a site’s current 

species. This has implications for survival probability, as well 

as for species that potentially immigrate to the site. The problem is explored using a set of heuristics for both of two policy 

objectives focusing on (1) the protection on current (native) 

species, and (2) all species, including immigrating species. The 

trade-offs between the protection of native species versus all 

species is illustrated. The study shows that the development of 

prediction models of future species distributions as the basis of 

decision rules can be crucial for ensuring the effectiveness of 

conservation plans. Finally, it is discussed how more adaptive 

strategies, that allow for the redirection of resources from protected sites to privately-owned sites, may increase the effectiveness of the conservation networks. Climate change induced 

shifts in the suitability of habitats for species may increase the 

value of such adaptive strategies, the benefit decreasing with 

increasing migration probabilities and species distribution 

dynamics.

196. Thorpe J. 2012. Adaptation to climate change in 

management of prairie grasslands. Saskatoon, Saskatchewan, 

Canada: Saskatchewan Research Council. SRC Publication 

No. 12855-1E12. 45 p.

Type: Report

Geographic Area: Canada

Compilers’ Keywords: migration, Saskatchewan

Summary: Options for adaptation to climate change in 

management of prairie grasslands were examined as part of 

Terrestrial Ecosystems component of the Prairies Regional 

Adaptation Collaborative (PRAC). Adaptation options were 

structured according to the “three Rs”: Create resistance 

CHAPTER 1 • CLIMATE CHANGE • STRATEGIES
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 71

in human social systems (e.g., management) in response to 

climate stimuli and their effects. Since management always 

occurs in the context of desired ecosystem conditions or natural resource management goals, it is instructive to examine 

particular goals and processes used by different organizations to fulfill their objectives. Such an examination allows 

for discussion of specific adaptation options as well as potential barriers and opportunities for implementation. Using 

this approach, this report presents a series of chapters on the 

following selected management systems: National Forests, 

National Parks, National Wildlife Refuges, Wild and Scenic 

Rivers, National Estuaries, and Marine Protected Areas. 

The information drawn from across these chapters is then 

analyzed to develop the key synthetic messages presented 

below.

Link: http://cfpub.epa.gov/ncea/cfm/recordisplay.

cfm?deid=180143

200. USDA FS. 2007. USDA Forest Service strategic plan FY 

2007-2012. Washington (DC): USDA Forest Service. FS-880. 

38 p.

Type: Government Document

Geographic Area: USA

Compilers’ Keywords: planning, forest management, climate change

Introduction: Managing the natural resources of the Nation’s 

forests and grasslands requires the complex integration of resource assessments, management actions, and cooperative 

partnerships. United States population growth and expanding urban centers have created a greater demand for goods, 

services, and amenities from the Nation’s private and public 

forests and grasslands. Given such changes, this section addresses core principles and issues central to delivering the 

Forest Service’s mission. Increasingly diverse urban populations are losing their awareness and knowledge of the natural 

systems on which they depend. The Forest Service, USDA, 

must connect with and educate these citizens to expand their 

understanding of the Links between people, the way they live, 

and the natural settings within which they live. With increasingly urbanized landscapes, emergency response—including 

fighting wildland fires—becomes more complex and challenging. Along with our partners, the Forest Service is committed to maintaining and fielding a safe, effective response 

organization that can be mobilized for managing wildland fires 

or other national emergencies. We are jointly committed to reducing the loss of life and property and maintaining landscape 

values. Together, we will invest in the personnel, training, and 

equipment and provide leadership commensurate with those 

responsibilities.

Link: http://www.fs.fed.us/publications/strategic/fs-spfy07-12.pdf

their development and control GHG emissions at relatively low 

costs. Opportunities are ample, but barriers are significant also. 

198. Turra A, Croquer A, Carranza A, Mansilla 

A, Areces AJ, Werlinger C, Martinez-Bayon C, 

Nassar CA, Plastino E, Schwindt E, Scarabino 

F, Chow F, Figueroa FL, Berchez F, Hall-Spencer 

JM, Soto LA, Buckeridge MS, Copertino MS, de 

Szechy MT, Ghilardi-Lopes NP, Horta P, Coutinho 

R, Fraschetti S, Leao ZM. 2013. Global environmental 

changes: setting priorities for Latin American coastal habitats. 

Global Change Biology 19:1965-1969.

Type: Journal 

Geographic Area: Central and South America

Keywords: benthic ecology, climate impacts, habitat mapping, long-term monitoring, marine biodiversity

Abstract: As the effects of the Global Climate Changes on 

the coastal regions of Central and South Americas advance, 

there is proportionally little research being made to understand 

such impacts. This commentary puts forward a series of propositions of strategies to improve performance of Central and 

South American science and policy making in order to cope 

with the future impacts of the Global Climate Changes in their 

coastal habitats.

Link: http://www.ncbi.nlm.nih.gov/pubmed/23504820

199. U.S. Climate Change Science Program 

2008. Preliminary review of adaptation options for climatesensitive ecosystems and resources. Washington (DC): U.S. 

Environmental Protection Agency. Synthesis and Assessment 

Product 4.4. 873 p.

Type: Government Document

Geographic Area: USA

Compilers’ Keywords: climate change, adaptation strategy, 

EPA, sensitive plants and animals

Summary: Climate variables are key determinants of 

geographic distributions and biophysical characteristics 

of ecosystems, communities, and species. Climate change 

is therefore having profound effects on species attributes, 

ecological interactions, and ecosystem processes. Because 

changes in the climate system will continue into the future 

regardless of emissions mitigation, strategies for protecting 

climate sensitive ecosystems through management will be 

increasingly important. While there will always be uncertainties associated with the future path of climate change, the 

response of ecosystems to climate impacts, and the effects of 

management, it is both possible and essential for adaptation 

to proceed using the best available science. This report provides a preliminary review of adaptation options for climatesensitive ecosystems and resources in the United States. The 

term “adaptation” in this document refers to adjustments 

CHAPTER 1 • CLIMATE CHANGE • STRATEGIES
72 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

(both rural and urban), grasslands, and their associated aquatic 

ecosystems (riparian systems, lakes, streams). The actions 

the Research Plan identifies will provide the scientific basis 

to sustain ecosystem health, adjust management for ecosystem services (“adaptation”) and increase carbon sequestration 

(“mitigation”), all under changing climate conditions. The fundamental research focus of the Research Plan is to increase 

scientists’ understanding of forest, woodland, and grassland 

ecosystems so that land managers can manage them in a 

way that sustains and provides ecosystem services for future 

generations.

Link: http://www.fs.fed.us/research/climate-change/

203. Valatin G. 2012. Additionality and climate change 

mitigation by the UK forest sector. Forestry 85:445-462.

Type: Journal 

Geographic Area: United Kingdom

Compilers’ Keywords: forest management, carbon 

accounting

Summary: Although widely considered to be a core aspect of 

quality assurance of climate change mitigation activities, additionality remains a source of much controversy in relation 

to carbon accounting and carbon markets. This article illuminates the multi-faceted nature of the concept and develops a 

taxonomy of different forms. It provides an overview of how 

additionality is currently applied in relation to both compliance and voluntary carbon markets, including tests used and 

underlying evidence base requirements. This draws upon and 

updates an earlier review commissioned to help inform development of a Woodland Carbon Code designed to underpin 

climate change mitigation activities in the UK by the forest 

sector. Sources of uncertainty and trade-offs in practical application of the concept are highlighted, and potential perverse 

incentives explored.

204. Wiens JA, Bachelet D. 2010. Matching the multiple scales of conservation with the multiple scales of climate 

change. Conservation Biology 24:51-62.

Type: Journal 

Geographic Area: Global

Keywords: climate models, climate change, conservation 

planning, downscaling, scale, short-grass prairie, The Nature 

Conservancy

Abstract: To anticipate the rapidly changing world resulting from global climate change, the projections of climate 

models must be incorporated into conservation. This requires 

that the scales of conservation be aligned with the scales of 

climate-change projections. We considered how conservation has incorporated spatial scale into protecting biodiversity, 

how the projections of climate-change models vary with scale, 

and how the two do or do not align. Conservation planners 

201. USDA FS. 2008. Forest service strategic framework for 

responding to climate change. Washington (DC): USDA Forest 

Service. Version 1.0. 21 p.

Type: Government Document

Geographic Area: USA

Compilers’ Keywords: management, forest resources, ecosystem services

Introduction: The Nation’s forests and grasslands provide 

clean water, scenic beauty, biodiversity, outdoor recreation, 

natural resource-based jobs, forest products, renewable energy and carbon sequestration. Climate change is one of the 

greatest challenges to sustainable management of forests and 

grasslands and to human well-being that we have ever faced, 

because rates of change will likely exceed many ecosystems’ 

capabilities to naturally adapt. Without fully integrating consideration of climate change impacts into planning and actions, 

the Forest Service can no longer fulfill its mission. The Forest 

Service has a unique opportunity and responsibility to sustain 

forests and grasslands in the United States and internationally. 

This responsibility includes: 1) stewardship of 193 million 

acres of national forests and grasslands, 2) partnerships with 

States, Tribes, and private landowners for assisting communities and owners of 430 million acres of private and Tribal 

forests, and with other federal agencies, 3) international cooperation, 4) research and development to provide science and 

management tools. These responsibilities make it imperative 

that we understand and be able to respond to the effects of climate change on the Nation’s forest and grassland resources. 

This document provides a strategic framework for the Forest 

Service to guide current and future actions to meet the challenge of climate change. It incorporates the actions included in 

Chief Gail Kimbell’s letter to the National Leadership Council 

of February 15, 2008.

Link: http://www.fs.fed.us/climatechange/message.shtml

202. USDA FS. 2010. Forest Service global change research 

strategy, 2009-2019 implementation plan. Washington (DC): 

USDA Forest Service, Research and Development. FS-948. 

72 p.

Type: Government Document

Geographic Area: USA

Compilers’ Keywords: climate change, forest management, 

assisted migration, USFS, mitigation

Summary: In keeping with the research goals of the U.S. 

Global Change Research Program, the climate change strategy of the USDA and the climate change framework of the 

Forest Service, this Forest Service Global Change Research 

Strategy, 2009-2019 Implementation Plan (hereafter called 

the Research Plan), was written by Forest Service Research 

and Development to help to define climate change policy and 

support best management practices for forests and woodlands 

CHAPTER 1 • CLIMATE CHANGE • STRATEGIES
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 73

use information about past and current ecological conditions 

at multiple scales to identify conservation targets and threats 

and guide conservation actions. Projections of climate change 

are also made at multiple scales, from global and regional 

circulation models to projections downscaled to local scales. 

These downscaled projections carry with them the uncertain-

ties associated with the broad-scale models from which they 

are derived; thus, their high resolution may be more apparent 

than real. Conservation at regional or global scales is about 

establishing priorities and influencing policy. At these scales, 

the coarseness and uncertainties of global and regional climate 

models may be less important than what they reveal about 

possible futures. At the ecoregional scale, the uncertainties as-

sociated with downscaling climate models become more criti-

cal because the distributions of conservation targets on which 

plans are founded may shift under future climates. At a local 

scale, variations in topography and land cover influence local 

climate, often overriding the projections of broad-scale climate 

models and increasing uncertainty. Despite the uncertainties, 

ecologists and conservationists must work with climatechange modelers to focus on the most likely projections. The 

future will be different from the past and full of surprises; judi-

cious use of model projections at appropriate scales may help 

us prepare.

Link: http://www.ncbi.nlm.nih.gov/pubmed/20121841

205. Williams SE, Shoo LP, Isaac JL, Hoffman AA, 

Langham G. 2008. Towards an integrated framework for 

assessing the vulnerability of species to climate change. PLoS 

Biology 6:2621-2626.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: species sensitivity, biodiversity con-

servation, tools, adaptive capacity

Introduction: We believe that progress in vulnerability assess-

ment relating to climate change could be hastened if a unified 

framework was available to coordinate the activities of disparate 

research disciplines. Specifically, what is needed is a complete 

working framework for assessing the vulnerability of species 

that explicitly links the various components of biotic vulnerabil-

ity; the regional and local factors determining exposure to cli-

matic change; the potential for both evolutionary and ecological 

responses, resilience, and active management to mediate the fi-

nal realised impacts; and the potential for feedback effects. Such 

a framework would be invaluable as it would integrate and guide 

thought, research programmes, and policy in the biodiversity/

climate change arena and allow significant gaps in knowledge to 

be clearly identified. To this end, we present a conceptual frame-

work that addresses these challenges.

Link: http://www.nccarf.edu.au/userfiles/williams_shoo_

etal2008_towards%20integrated%20framework.pdf
206. Bagne KE, Friggens MM, Finch DM. 2011. A 

system for assessing vulnerability of species (SAVS) to climate 

change. Fort Collins (CO): USDA Forest Service, Rocky 

Mountain Research Station. General Technical Report RMRSGTR-257. 28 p.

Type: Government Document

Geographic Area: USA

Keywords: climate change, vulnerability, vertebrate species, 

conservation, resource management

Abstract: Sustained conservation of species requires integration of future climate change effects, but few tools exist to assist 

managers. The System for Assessing Vulnerability of Species 

(SAVS) identifies the relative vulnerability or resilience of vertebrate species to climate change. Designed for managers, the 

SAVS is an easily applied tool that uses a questionnaire of 22 

predictive criteria to create vulnerability scores. The user scores 

species’ attributes relating to potential vulnerability or resilience 

associated with projections for their region. Six scores are produced: an overall score denoting level of vulnerability or resilience, four categorical scores (habitat, physiology, phenology, 

and biotic interactions) indicating source of vulnerability, and an 

uncertainty score, which reflects user confidence in the predicted 

response. The SAVS provides a framework for integrating new 

information into the climate change assessment process.

Link: http://www.treesearch.fs.fed.us/pubs/37850

207. Bagne KE, Friggens MM, Finch DM. 2011.

SAVS: a system for assessing vulnerability of species.

Albuquerque (NM): USDA Forest Service, Rocky 

Mountain Research Station. (URL accessed 07 November 

2015)

Type: Web Page

Geographic Area: USA

Compilers’ Keywords: decision support system, terrestrial 

vertebrates, resource management, climate change, tool, USFS 

Compilers’ Summary: The Rocky Mountain Research Station 

(USFS) has developed a System for Assessing Vulnerability of 

Species (SAVS) that quantifies the relative impact of expected 

climate change effects for terrestrial vertebrate species. The 

SAVS uses 22 criteria related to expected response or vulnerability of species in a questionnaire to provide a framework for 

assessing vulnerability to climate change. The questionnaire is 

completed using information gathered from published materials, 

personal knowledge, or expert consultation. The SAVS Climate 

Change Tool focuses solely on the effects of climate change for 

terrestrial vertebrate species. This tool aids managers by identifying specific traits and issues related to individual species 

General (1–36)—editorials, opinions, 

reviews, trends, surveys, summaries

Research (37–135)—climate change impacts, 

projections, paleoecology

Strategies (136–205)—policies, adaptation 

options, frameworks, mitigations

Resources (206–214)—tools, websites, 

software

Climate Change 

CHAPTER 1 • CLIMATE CHANGE • RESOURCES

#. Author(s), Date. Title. Source.

Type: 

Geographic Area: 

Keywords: 

Description:

Link:

location of study

keywords written by the 

author(s) or provided by the 

compilers

ID number and

information

abstract, summary, or 

portions of the introduction 

or conclusion written by the 

author(s) or provided by the 

compilers

book, book section, conference 

paper, dissertation, government 

document, journal article, 

newsletter, presentation, report, 

thesis or web page

internet address of reference

Each reference contains the following:

74 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 75

climate change considerations into project planning as 

well as for developing concrete adaptation options for land 

managers. We designed CPST as part of the Westwide Climate 

Initiative project, which seeks to develop adaptation options for 

addressing climate change through science/management partnerships. The CPST lists projected climate trends for the target 

region and questions to be considered when designing projects 

in different resource areas. The objective is to explore options 

for ameliorating the effects of climate on resource management 

projects. To pilot the CPST, we interviewed 13 staff members 

and line officers of the U.S. Forest Service and Bureau of Land 

Management in the Sierra Nevada region of California. We 

found that a major value of the CPST was the process—with 

the activity of conducting the questionnaire being as important 

as the answers received from the staff. The CPST also serves 

as a priority-setting tool, allowing managers to consider effects 

of different actions. Finally, the CPST helps to reduce uncertainty by identifying the range of impacts that both climatic 

changes and management actions may have on resources. The 

CPST could also be modified to devise mitigation options for 

resource managers.

Link: http://www.treesearch.fs.fed.us/pubs/40319

210. Phillips NC, Anderson RP, Schapire RE. 2006.

MaxEnt software for species habitat modeling. (URL accessed 07 

November 2015). 

Type: Web Page

Geographic Area: Global

Compilers’ Keywords: MaxEnt, climate change, software, 

decision support tool, climate projections, modeling, species 

distribution

Compilers’ Summary: MaxEnt is a program for modelling 

species distributions from presence-only species records. The 

software has been used to estimate species distributions under 

climate change scenarios, in an effort to evaluate management 

options such as assisted migration (see Vitt et al. 2010). 

Link: http://www.cs.princeton.edu/~schapire/maxent/

211. Potter KM, Crane BS. 2010. Forest tree genetic risk 

assessment systems: a tool for conservation decision-making in 

changing times. Atlanta (GA): USDA Forest Service. 

Type: Computer Program

Geographic Area: USA

Compilers’ Keywords: user guide, climate change, tool, 

ForGRAS, sensitivity, forest management

Overview: Changing climate conditions and increasing 

pest and pathogen infestations will increase the likelihood 

that forest trees could experience population-level extirpation or species-level extinction during the next century. 

Funds, however, will be limited for management and gene 

conservation efforts to preserve forest tree genetic diversity. 

vulnerabilities. Scores generated by completing a questionnaire 

are meant to be used to inform management planning. A comprehensive discussion of the development and application of this 

tool will be available from the RMRS General Technical Report 

[A System for Assessing Vulnerability of Species (SAVS) to 

Climate Change]. Information regarding specific criteria for 

each question are provided as information pop-ups with the 

scoring template. Users may also find it helpful to review assessments conducted for species in New Mexico and Arizona 

and legacy project briefs for Fort Huachuca, AZ and Barry M 

Goldwater Range, AZ. 

Link: http://www.fs.fed.us/rm/grassland-shrubland-desert/

products/species-vulnerability/

208. Devine W, Aubry C, Miller J, Potter KM, 

Bower AD. 2012. Climate change and forest trees in the 

Pacific Northwest: guide to vulnerability assessment methodology. Olympia (WA): USDA Forest Service, Pacific Northwest 

Region. 49 p.

Type: Government Document

Geographic Area: Pacific Northwest USA

Compilers’ Keywords: tool, forest management, tree species, 

risk assessment, model, USFS, ForGRAS

Introduction: This guide provides a step-by-step description 

of the methodology used to apply the Forest Tree Genetic Risk 

Assessment System (ForGRAS) to the tree species of the Pacific 

Northwest in a recent climate change vulnerability assessment. 

We describe our modified version of the ForGRAS model, and 

we review the model’s basic principles and operation. We also 

discuss further adjustments to the model that managers may find 

useful if applying it to other regions or to fit other objectives. 

Chapter 2 of this guide describes the overall methodology of the 

vulnerability assessment; chapters 3 through 7 describe each of 

the five risk factors that we selected for the Pacific Northwest 

vulnerability assessment. Appendices provide step-by-step procedures used to prepare the model input data.

Link: http://www.treesearch.fs.fed.us/pubs/44236

209. Morelli TL, Yeh S, Smith NM, Hennessy MB, 

Millar CI. 2012. Climate project screening tool: an aid for 

climate change adaptation. Albany (CA): USDA Forest Service, 

Pacific Southwest Research Station. Research Paper PSWRP-263. 29 p.

Type: Government Document

Geographic Area: Western USA

Keywords: climate change, adaptation, land management, 

mitigation, Sierra Nevada

Abstract: To address the impacts of climate change, land 

managers need techniques for incorporating adaptation 

into ongoing or impending projects. We present a new tool, 

the Climate Project Screening Tool (CPST), for integrating 

CHAPTER 1 • CLIMATE CHANGE • RESOURCES
76 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Compilers’ Keywords: decision support system, online tool, 

genetic adaptation, genetic degradation, forest management, 

Pacific Northwest, Southeastern United States, tree species, climate change

Summary: A variety of threats, most importantly climate 

change and insect and disease infestation, will increase the 

risk that forest trees could experience population-level or 

species-level extinction. Species, however, differ in important traits such as life-history strategies and population 

dynamics, which could drive widely varying responses to 

potential threats. Determining how to prioritize species 

for management and conservation activities in the face of 

these threats will pose a particular challenge in speciesrich regions. To address this challenge, a cooperating scientist with the Southern Research Station’s Eastern Forest 

Environmental Threat Assessment Center has developed a 

framework that allows managers to assess the relative risk 

of genetic degradation to forest trees affected by multiple 

threats. This assessment framework serves as a tool for 

planning management activities and conservation efforts, 

for evaluating species’ genetic resources, and for detecting vulnerabilities. It has the advantage of accounting for 

multiple threats that may result in the most severe genetic 

impacts. Only by considering population-level extirpation 

as a synergistic process of external threats and intrinsic biological traits will we be able to make predictions of risk that 

approximate reality for most species. Known as the Forest 

Tree Genetic Risk Assessment System (ForGRAS), the 

framework has been used by the Forest Service Southern 

and Pacific Northwest Regions to identify species at risk 

as a step towards developing management plans. The flexibility of this approach allows for its application at multiple 

scales and across any area for which data exist on the population dynamics and distribution of the species of interest. 

Link: http://www.forestthreats.org/research/projects/project-summaries/genetic-risk-assessment-system

213. Prasad AM, Iverson LR, Matthews S, Peters 

M. 2007. A climate change atlas for 134 forest tree species of the 

eastern United States [database]. Delaware (OH): USDA 

Forest Service, Northern Research Station. (URL accessed 07 November 2015)

Type: Web Page

Geographic Area: Eastern USA

Compilers’ Keywords: sensitivity analysis, models, GCM, 

decision support system, forest management

Summary: We assess the current status (2000) and potential future status (2100) following climate change, of 134 

tree species in the eastern United States (east of 100th meridian). We use U.S. Forest Service inventory data with 38 

environmental variables to generate models of current suitable habitat for each species. We then change the climate 

The Forest Tree Genetic Risk Assessment System provides a 

framework for users to rank the relative risk of genetic degradation for multiple forest tree species. We have applied 

this framework for the species-rich forests of the southern 

Appalachian Mountains of the southeastern United States. 

Species differ in their physiological tolerances, life history strategies, probabilities of population extinctions and 

colonizations, and dispersal abilities; these individualistic traits likely underlie the high variability in strength of 

climate response across wild species, even among those 

subjected to similar climatic trends. In the face of these 

changes, an important goal will be to safeguard existing 

adaptedness and create conducive conditions for future 

evolution, with a focus on the conservation of variability 

in adaptive traits. In the absence of good information about 

the genetic composition of species, or even an understanding of the relevant adaptive traits, several researchers have 

suggested using ecological and life-history traits to rank 

the predisposition of species to climate change and other 

threats, for conservation planning, for the evaluation of 

species’ genetic resources, and for the early detection of 

vulnerability. The Forest Tree Genetic Risk Assessment 

System gives each species a rating for risk factors relating to (1) its intrinsic attributes, such as population structure, fecundity and seed dispersal mechanism, that may 

increase its vulnerability in the face of change, and (2) the 

external threats to its genetic integrity, including changing climate and insect and disease threats. Species may 

be additionally rated for a set of conservation modifiers, 

such as listed status and endemism. The factor index values are weighted and summed to give risk ratings for the 

species within a given region, which are ranked according 

to their overall susceptibility to genetic degradation. This 

system has the advantage of accounting for the interactions among threats that may result in the most severe impacts associated with climate change. Only by considering 

extinction as a synergistic process of external threats and 

intrinsic biological traits will it be possible to make predictions of risk that approximate reality for most species, 

and therefore to increase the likelihood that conservation 

efforts will be effective. Additionally, the flexibility of the 

Forest Tree Genetic Risk Assessment System allows for 

its application at multiple scales and across any area for 

which the relevant data exist for the species of interest.

Link: http://www.forestthreats.org/research/projects/project-summaries/genetic-risk-assessment-system

212. Potter KM, Crane BS. 2010. Forest tree genetic 

risk assessment system (ForGRAS). North Carolina State 

University and USDA Forest Service. (URL accessed 07 

November 2015)

Type: Web Page

Geographic Area: USA

CHAPTER 1 • CLIMATE CHANGE • RESOURCES
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 77

according to three General Circulation Model (GCM) climate models (HADCM3, PCM & GFDL—see 3-GCMs 

for details) and two emissions scenarios (A1FI (Hi) = little 

conservation efforts to mitigate CO2 emissions, B1 (Lo) = 

significant conservation effort), and model the potential 

future species habitats. These two emissions scenarios 

bracket most of the emission futures as outlined by the 

Intergovernmental Panel on Climate Change’s (IPCC) 

evaluation of emission scenarios (Nakicenovic et al. 2000), 

and end the 21st century at roughly double (550 ppm-B1) 

and triple (970 ppm-A1fi) the pre-industrial levels of CO2.

Link: http://www.nrs.fs.fed.us/atlas/tree

214. Regniere J, Saint-Amant R. 2008. BioSIM 9 - 

user’s manual. Natural Resources Canada, Canadian Forest 

Service, Laurentian Forestry Centre. Information Report 

LAU-X-134. 76 p.

Type: Government Document

Geographic Area: Canada

Compilers’ Keywords: software, simulation model, pest management, forecasting, climate change, seasonal biology

Introduction: BioSIM is a software tool designed to assist 

in the application of temperature-driven simulation models in pest management. It can also be used as a tool in the 

development and analysis of such models for the purposes 

of scientific investigation. However, BioSIM’s main purpose is to generate forecasts of features or “events” in the 

seasonal biology of pests or their host plants. Forecasts 

are made by simulation models provided by the system 

and are based on regional air temperature and precipitation 

interpolated from nearby weather stations, adjusted for elevation and location differentials with regional gradients. 

Link: http://www.publications.gc.ca/collections/collection_2008/nrcan/Fo113-3-134E.pdf

CHAPTER 1 • CLIMATE CHANGE • RESOURCES
78 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

215. Arthur W. 1987. The niche in competition and evolution. New York (NY): John Wiley and Sons. 175 p. 

Type: Book

Geographic Area: Global

Compilers’ Keywords: population ecology, plant community ecology

Summary: Using the concept of the niche (and multipleniche) this book attempts to synthesize two widelyinvestigated topics: polymorphism and between-species 

competition; the author’s view being that multiple-niche 

coexistence and multiple-niche polymorphism together 

represent a common means of coexistence of competitors. 

Experimental detail and fieldwork are frequently cited 

in order to create a common point of reference for both 

population ecologists and geneticists. Concluding chapters 

discuss other important principles and hypotheses in 

evolution which are related to the niche concept.

216. Breukers A, van Asseldonk M, Bremmer J, 

Beekman V. 2012. Understanding growers’ decisions to 

manage invasive pathogens at the farm level. Phytopathology 

102:609-619.

Type: Journal

Geographic Area: Global

Compilers’ Keywords: risk management, nursery practices, 

climate change, invasive species, pests, disease control and 

management

Abstract: Globalization causes plant production systems to 

be increasingly threatened by invasive pests and pathogens. 

Much research is devoted to support management of these 

risks. Yet, the role of growers’ perceptions and behavior in 

risk management has remained insufficiently analyzed. This 

article aims to fill this gap by addressing risk management of 

invasive pathogens from a sociopsychological perspective. 

An analytical framework based on the Theory of Planned 

Behavior was used to explain growers’ decisions on voluntary risk management measures. Survey information from 

303 Dutch horticultural growers was statistically analyzed, 

including regression and cluster analysis. It appeared that 

growers were generally willing to apply risk management 

measures, and that poor risk management was mainly due to 

perceived barriers, such as high costs and doubts regarding 

efficacy of management measures. The management measures applied varied considerably among growers, depending on production sector and farm-specific circumstances. 

General (215–232)—reviews, surveys, 

summaries, theory, foundations, paradigms

Research (233–285)—propagation, outplanting, invasion

Strategies (286–326)—frameworks, approach, decision support

Resources (327–338)—native plant materials, 

guidelines, handbooks

Conservation and 

Restoration

CHAPTER 2 • CONSERVATION AND RESTORATION • GENERAL

#. Author(s), Date. Title. Source.

Type: 

Geographic Area: 

Keywords: 

Description:

Link:

location of study

keywords written by the 

author(s) or provided by the 

compilers

ID number and

information

abstract, summary, or 

portions of the introduction 

or conclusion written by the 

author(s) or provided by the 

compilers

book, book section, conference 

paper, dissertation, government 

document, journal article, 

newsletter, presentation, report, 

thesis or web page

internet address of reference

Each reference contains the following:
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

218. Campbell FT, Schlarbaum SE. 2014. Fading 

forests III: American forests, what choice will we make?

Arlington (VA): The Nature Conservancy. 167 p.

Type: Report

Geographic Area: USA

Compilers’ Keywords: forest management, invasive species, pest management, restoration, invasive pathogens, policy

Preamble (4th paragraph): Fading Forests III is the last 

publication of a trilogy on pests, policy, and forest health that 

we began over 20 years ago. This publication, however, is not 

intended to be a requiem. Instead, we hope that Fading Forests 

III will be received as a call to greater effort, focused on a limited agenda of actions that can ensure our forests’ future.

Link: http://www.nature.org/ourinitiatives/habitats/forests/

fading-forests-3-complete-report.pdf

219. Choi YD. 2007. Restoration ecology to the future: a 

call for new paradigm. Restoration Ecology 15:351-353.

Type: Journal 

Geographic Area: Global

Keywords: ecology, future, paradigm, rehabilitation, restoration, sustainability

Abstract: The discipline of restoration ecology has grown 

remarkably in the past decades, providing new ideas and opportunities for conserving biological diversity, managing ecosystems, and testing ecological theories. On the other side, its 

past-oriented, static, and idealistic approach has been criticized 

for subjectivity in determining restoration goals, inapplicability to dynamic ecosystems, and inability for restoring certain irreversible losses. Moreover, unpredictable sustainability of the 

restored ecosystems, which were modeled after its historical 

fidelity, adds our skepticism under the changing environment. 

This paper calls for a new paradigm of ecological restoration to 

the future. A future-oriented restoration should (1) establish the 

ecosystems that are able to sustain in the future, not the past, 

environment; (2) have multiple alternative goals and trajectories for unpredictable endpoints; (3) focus on rehabilitation of 

ecosystem functions rather than recomposition of species or 

cosmetics of landscape surface; and (4) acknowledge its identity as a ‘‘value-laden’’ applied science within economically 

and socially acceptable framework. Applicability of ecological 

theories to restoration practice is also discussed in this paper.

Link: http://www.globalrestorationnetwork.org/uploads/

files/LiteratureAttachments/564_restoration-ecology-to-thefuture—a-call-for-a-new-paradigm.pdf

220. Handel SN. 2011. Past, present, future. Ecological 

Restoration 29:203-205.

Type: Journal 

Geographic Area: Global

79

Growers’ risk perception was found to play a role in their 

risk management, although the causal relation remained unclear. These results underscore the need to apply a holistic 

perspective to farm level management of invasive pathogen 

risk, considering the entire package of management measures and accounting for sector- and farm-specific circumstances. Moreover, they demonstrate that invasive pathogen 

risk management can benefit from a multidisciplinary approach that incorporates growers’ perceptions and behavior.

Link: http://www.ncbi.nlm.nih.gov/pubmed/22397409

217. Burton PJ, Macdonald SE. 2011. The restorative 

imperative: challenges, objectives and approaches to restoring 

naturalness in forests. Silva Fennica 45:843-863.

Type: Journal 

Geographic Area: Global

Keywords: afforestation, disturbance regime, ecological restoration, forest rehabilitation, native species, reclamation

Abstract: Many of the world’s forests are not primeval; forest restoration aims to reverse alterations caused by human 

use. Forest restoration (including reforestation and forest 

rehabilitation) is widely researched and practiced around 

the globe. A review of recent literature reveals some common themes concerning forest restoration motivations and 

methods. In some parts of the world, forest restoration aims 

mainly to re-establish trees required for timber or fuel wood; 

such work emphasizes the propagation, establishment and 

growth of trees, and equates with the traditional discipline of 

silviculture. Elsewhere, a recent focus on biocentric values 

adopts the goal of supporting full complements of indigenous trees and other species. Such ecosystem based restoration approaches consider natural templates and a wide array 

of attributes and processes, but there remains an emphasis 

on trees and plant species composition. Efforts to restore 

natural processes such as nutrient cycling, succession, and 

natural disturbances seem limited, except for the use of fire, 

which has seen widespread adoption in some regions. The 

inherent challenges in restoring “naturalness” include high 

temporal and spatial heterogeneity in forest conditions and 

natural disturbances, the long history of human influence on 

forests in many regions of the world, and uncertainty about 

future climate and disturbance regimes. Although fixed templates may be inappropriate, we still have a reasonably clear 

idea of the incremental steps required to make forests more 

natural. Because most locations can support many alternative configurations of natural vegetation, the restoration of 

forest naturalness necessarily involves the setting of priorities and strategic directions in the context of human values 

and objectives, as informed by our best understanding of 

ecosystem structure and function now and in the future.

Link: http://www.silvafennica.fi/pdf/article74.pdf

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80

223. Landis TD. 2011. The target plant concept—a history and brief overview. In: Riley LE, Haase DL, Pinto JR, 

editors. Forest and Conservation Nursery Associations—2010. 

Fort Collins (CO): USDA Forest Service, Rocky Mountain 

Research Station. Proceedings RMRS-P-65. p 61-66.

Type: Government Document

Geographic Area: Global

Keywords: nursery, reforestation, restoration, seedling, native 

plant

Abstract: The target plant concept originated with morphological classification of conifer nursery stock in the 1930s, and 

the concept was enhanced through physiological research and 

seedling testing towards the end of the century. Morphological 

grading standards such as shoot height, stem diameter, and root 

mass are the most common use of the target plant concept, and 

some physiological grading standards are also being operationally implemented by nursery workers and seedling users. Since 

2000, the concept has been expanded to include all types of 

plant materials, including seeds, cuttings, or wildlings, as well 

as traditional nursery stock. Because these native plant materials are being outplanted on harsh, severely disturbed sites, 

this more comprehensive native plant materials concept also 

involves environmental conditions on the project site. 

Link: http://www.treesearch.fs.fed.us/pubs/40170

224. Norton DA, Forbes A. 2013. Can exotic pine trees 

assist in restoration? Applied Vegetation Science 16:169-170.

Type: Journal 

Geographic Area: Chile

Compilers’ Keywords: conservation, exotic species, 

regeneration

Abstract: Invasive exotic woody species, including conifers escaped from plantations, are usually regarded as serious 

threats to native biodiversity. Becerra & Montenegro (Applied 

Vegetation Science, 16, 2013, 2) present an interesting example of invasive Pinus radiata facilitating native woody regeneration in semi-arid central Chile. However, the positive value 

of invasive conifers for restoration and conservation will not 

necessarily apply in all situations.

Link: http://onlinelibrary.wiley.com/doi/10.1111/avsc.12014/

full

225. Oliet JA, Jacobs DF. 2012. Restoring forests: advances in techniques and theory. New Forests 2012:1-7.

Type: Journal

Geographic Area: Mediterranean

Compilers’ Keywords: genetics, disturbance, climate 

change, ecological function, review

Introduction: Much of the past science of forest regeneration 

emphasized reforestation following timber harvest for industrial 

Compilers’ Keywords: restoration, climate change, editorial

Closing Remarks: Importance of restoration ecology must 

enlarge and quicken if we are going to keep pace with environmental change and deliver the ecological services upon which 

we all depend. We can only hope that this journal is carried 

through. 

Link: http://er.uwpress.org/content/29/3/203.full.pdf

221. Jones TA. 2009. Conservation biology and plant 

breeding: special considerations for the development of native 

plant materials for use in restoration. Ecological Restoration 

27:8-11.

Type: Journal 

Geographic Area: USA

Compilers’ Keywords: nursery, genetic diversity, evolution, 

practices

Introduction: The development of native plant materials for 

restoration demands that close attention be paid to the expectations of the specialized customer base of ecological restorationists. Restoration practitioners’ expectations are often built 

on assumptions and values reflective of the relatively new 

discipline of conservation biology. Unlike most scientific disciplines, conservation biology is unashamedly value driven. Its 

basic thrust is that we must maintain natural patterns of genetic 

diversity at many levels and thus preserve options for future 

evolution.

Link: http://sfc.smallfarmcentral.com/dynamic_content/uploadfiles/152/jones-conservation%20biology.pdf

222. Jones TA, Young SA. 2005. Native seeds in commerce: more frequently asked questions. Native Plants Journal 

6:286-293.

Type: Journal

Geographic Area: Global

Keywords: seed prices, release, cultivar, pre-variety germplasm, genetic shift, pure live seed, seed certification, variety, 

germplasm, plant variety protection

Abstract: To make intelligent choices in the marketplace, 

customers should have a working understanding of 1) the diversity of products that are available in the seed trade; 2) how 

they move in the market; 3) how they are regulated for purity 

and germination; 4) how they are certified to assure genetic 

identity; and 5) how they are legally protected as intellectual 

property. Options for seed certification have greatly increased 

over the last several years. Two complementary certification mechanisms are currently being used to deliver native 

plant materials, one facilitating the traditional cultivars and 

the other directed toward the novel pre-variety germplasms. 

Both accommodate natural and genetically manipulated plant 

materials.

CHAPTER 2 • CONSERVATION AND RESTORATION • GENERAL

USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015
81

227. Ratnam W, Rajora OP, Finkeldey R, 

Aravanopoulos F, Bouvet J-M, Vaillancourt RE, 

Kanashiro M, Fady B, Tomita M, Vinson C. 2014.

Genetic effects of forest management practices: global synthesis 

and perspectives. Forest Ecology and Management 333:52-65.

Type: Journal 

Geographic Area: Global

Keywords: forest management practices, timber production, 

genetic diversity, population structure, mating system, temperate, boreal and tropical forests

Abstract: Understanding the genetic impacts of forest 

management practices is crucial for conservation and management of forest genetic resources. Forest management 

practices based on selective and clear cut systems followed 

by natural or artificial regeneration can impact population 

structure and mating patterns, thus gene flow and genetic 

diversity. Survival and productivity of both tree and nontree species can be compromised or, possibly, enhanced. 

The extent of genetic impacts depends on the management 

system applied, stand structure as well as species’ distribution, demography, biological attributes and ecology. The 

impact of management practices is reviewed and synthesized for temperate, boreal and tropical forests based on experimental and simulation studies. In addition, the effects of 

genetically improved planting materials and establishment 

of large scale plantations on natural forests are examined. 

Recommendations are made for genetically sustainable forest management practices.

Link: http://www.sciencedirect.com/science/article/pii/

S0378112714003697

228. Simberloff D. 1980. A succession of paradigms 

in ecology: essentialism to materialism and probabilism.

Synthese 43:3-39.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: evolution, genetics, Darwin, review, 

foundational ecology

Compilers’ Summary: In this essay, Simberloff discusses 

the major paradigms in ecology beginning with the materialistic revolution in evolution and genetics and ending with 

the ecosystem paradigm. Ecology’s first paradigm that plant 

community is a superorganism (circa 1905 by Clements) 

was challenged by several ecologists, because succession 

does not always lead to the same climax. Simberloff claims 

that Clements’ superorganism is not obsolete, however, but 

has transformed into a belief that the study of ecosystems 

should be approached in a holistic manner. Much of the information provided by Simberloff has shaped the way we 

study, conserve and restore ecosystems. 

purposes. In recent decades, a pronounced evolution has occurred 

whereby the focal point of many of today’s forest regeneration 

scientists has shifted prominently toward restoration of harsh, 

degraded environments. Forest restoration is a highly complex 

process that requires consistent shifts to adapt to emerging circumstances. This is a special issue of 27 review articles covering 

a wide range of diverse issues in forest restoration. 

Link: http://www.researchgate.net/publication/236148435_

Restoring_forests_advances_in_techniques_and_theory/file/

e0b4951665abeca599.pdf

226. Rajora OP, Mosseler A. 2001. Challenges and 

opportunities for conservation of forest genetic resources.

Euphytica 118:197-212.

Type: Journal 

Geographic Area: Canada

Keywords: adaptive traits, gene and genetic resource conservation, genetic and ecological restoration, genetic diversity 

and population genetic parameters, minimum viable population size, molecular markers, sustainable forest management

Abstract: Increased use of forest resources and a shrinking 

forestland base threaten the sustainability of forest genetic 

resources and highlight the importance of conservation and 

sustainable management of these resources. As forest trees are 

normally the keystone species of forest ecosystems, their continued existence is essential for many floral and faunal associations of these ecosystems. Major concepts, challenges and 

opportunities for conservation of forest genetic resources are 

briefly discussed in this paper. The major challenges include 

population decline and population structure changes due to 

forest removal and conversion of forest land to other uses, forest fragmentation, forestry practices, climate change, disease 

conditions, introduced pests, atmospheric pollution, and introgressive hybridization. Developing scientifically sound conservation strategies, maintaining minimum viable population 

sizes, and deployment of genetically engineered organisms 

represent other important challenges in conservation. The usefulness of various biochemical and molecular genetic markers, 

adaptive traits, and genetic diversity measures for developing 

conservation strategies for in situ and ex situ genetic resource 

conservation are also discussed. Major opportunities for conservation of forest genetic resources include: use of molecular 

genetic markers and adaptive traits for developing conservation strategies; in situ conservation through natural reserves, 

protected areas, and sustainable forest management practices; 

ex situ conservation through germplasm banks, common garden archives, seed banks, DNA banks, and tissue culture and 

cryopreservation; incorporation of disease, pest, and stress 

tolerance traits through genetic transformation; plantation forestry; and ecological restoration of rare or declining tree species and populations. Forest genetic resource conservation and 

resource use should be considered complementary rather than 

contradictory to each other.

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82

the biophysical dimensions. Social considerations can trump 

biophysical factors.

Link: http://www.treesearch.fs.fed.us/pubs/46976

230. Stanturf JA, Palik BJ, Williams MI, Dumroese 

RK, Madsen P. 2014. Forest restoration paradigms. Journal 

of Sustainable Forestry 33:S161-S194.

Type: Journal 

Geographic Area: Global

Keywords: reconstruction, rehabilitation, reclamation, novel 

ecosystems, intervention ecology

Abstract: An estimated 2 billion ha of forests are degraded 

globally and global change suggests even greater need for forest restoration. Four forest restoration paradigms are identified 

and discussed: revegetation, ecological restoration, functional 

restoration, and forest landscape restoration. Restoration is 

examined in terms of a degraded starting point and an ending point of an idealized natural forest. Global change, climate 

variability, biotechnology, and synthetic biology pose significant challenges to current restoration paradigms, underscoring 

the importance of clearly defined goals focused on functional 

ecosystems. Public debate is needed on acceptable goals; one 

role for science is to inform and help frame the debate and 

describe feasibility and probable consequences.

Link: http://www.treesearch.fs.fed.us/pubs/45775

231. Watrud E, Zensen F, Darbyshire R. 2012.

Laws affecting reforestation on USDA Forest Service lands. Tree 

Planters’ Notes 55:39-42.

Type: Government Document

Geographic Area: USA

Compilers’ Keywords: USFS, policy, restoration, management, national forests, silviculture

Abstract: Many laws affect reforestation practices on U.S. 

Department of Agriculture (USDA), Forest Service lands. 

This article summarizes several acts that have had important 

influences on Federal reforestation. In particular, we delve into 

The Knutson-Vandenberg Act of 1930 and the National Forest 

Management Act of 1976, which have had the largest effect on 

reforestation of the national forests.

Link: http://www.rngr.net/publications/tpn/55-2/laws-affecting-reforestation-on-usda-forest-service-lands/at_download/

file

232. Watt AS. 1947. Pattern and process in the plant community. Journal of Ecology 35:1-22.

Type: Journal 

Geographic Area: Global

229. Stanturf JA, Palik BJ, Dumroese RK. 2014.

Contemporary forest restoration: a review emphasizing function. Forest Ecology and Management 331:292-323.

Type: Journal 

Geographic Area: Global

Keywords: reconstruction, rehabilitation, reclamation, replacement, ecological restoration, forest landscape restoration

Abstract: The forest restoration challenge (globally 2 billion ha) and the prospect of changing climate with increasing 

frequency of extreme events argues for approaching restoration from a functional and landscape perspective. Because the 

practice of restoration utilizes many techniques common to 

silviculture, no clear line separates ordinary forestry practices 

from restoration. The distinction may be that extraordinary 

activities are required in the face of degraded, damaged, or 

destroyed ecosystems. Restoration is driven by the desire to 

increase sustainability of ecosystems and their services and 

restoration is likely to have multiple goals arising from the 

motivations of those involved. The process of setting restoration objectives translates vague goals into feasible, measurable 

targets and ultimately actions on the ground. Our objective 

for this review is to synthesize the science underpinning contemporary approaches to forest restoration practice. We focus 

on methods and present them within a coherent terminology 

of four restoration strategies: rehabilitation, reconstruction, 

reclamation, and replacement. While not a consensus terminology, these terms have a logical foundation. Rehabilitation 

restores desired species composition, structure, or processes 

to a degraded ecosystem. Reconstruction restores native plant 

communities on land recently in other resource uses, such as 

agriculture. Reclamation restores severely degraded land generally devoid of vegetation, often the result of resource extraction, such as mining. Replacement of species (or their locallyadapted genotypes) with new species (or new genotypes) is a 

response to climate change. Restoration methods are presented 

as available tools; because adding vegetation is an effective 

restoration technique, the discussion of methods begins with 

a description of available plant materials. We then discuss altering composition under different initial overstory conditions, 

including deployment methods depending upon whether or not 

an overstory is present, how much of the landscape will be 

restored, and the complexity of the planting design. We present 

some major approaches for altering structure in degraded forest stands, and describe approaches for restoration of two key 

ecosystem processes, fire and flooding. Although we consider 

stand-level designs, what we describe is mostly scalable to the 

landscape-level. No restoration project is undertaken in a social 

vacuum; even stand-level restoration occurs within a system 

of governance that regulates relationships among key agents. 

Gathering information and understanding the social dimensions of a restoration project is as necessary as understanding 

CHAPTER 2 • CONSERVATION AND RESTORATION • GENERAL

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83

the field of inquiry is limited to the plant community, divorced 

from its context in the sere; all reference to relics from its antecedents and to invaders from the next state is omitted. I am 

assuming essential uniformity in the fundamental factors of the 

habitat and essential stability of the community over a reasonable period of time.

Link: http://faculty.washington.edu/edford/Watt_1947.pdf

Compilers’ Keywords: ecology, patch dynamics, structure

Introduction: The plant community may be described from 

two points of view, for diagnosis and classification, and as 

a working mechanism. My primary concern is with the second of these. But in as much as the two aspects are not mutually exclusive, a contribution to our understanding of how 

a community is put together, and how it works, may contain 

something of value in description for diagnosis. It is now half 

a century since the study of ecology was injected with the dynamic concept, yet in the vast output of literature stimulated 

by it there is no record of an attempt to apply dynamic principles to the elucidation of the plant community itself and to 

formulate laws according to which it maintains and regenerates itself. Pavillard’s assessment of the dynamic behaviour of 

species comes very near it, but is essentially concerned with 

the ‘influence (direct or indirect) of the species on the natural 

evolution of plant communities. As things are, the current descriptions of plant communities provide information of some, 

but not critical, value to an understanding of them; how the 

individuals and the species are put together, what determines 

their relative proportions and their spatial and temporal relations to each other, are for the most part unknown. It is true 

that certain recent statistical work is stretching out towards that 

end, but the application of statistical technique, the formulation 

of laws and their expression in mathematical terms, will be 

facilitated if an acceptable qualitative statement of the nature 

of the relations between the components of the community is 

first presented. Such a statement is now made based on the 

study of seven communities in greater or less detail, for data 

of the kind required are seldom recorded. The ultimate parts 

of the community are the individual plants, but a description 

of it in terms of the characters of these units and their spatial 

relations to each other is impracticable at the individual level. 

It is, however, feasible in terms of the aggregates of individuals and of species which form different kinds of patches; these 

patches form a mosaic and together constitute the community. 

Recognition of the patch is fundamental to an understanding 

of structure as analysed here. In the subsequent analysis evidence is adduced to show that the patches (or phases, as I am 

calling them) are dynamically related to each other. Out of this 

arises that orderly change which accounts for the persistence 

of the pattern in the plant community. But there are also departures from this inherent tendency to orderliness caused by 

fortuitous obstacles to the normal time sequence. At any given 

time, therefore, structure is the result of causes which make for 

order and those that tend to upset it. Both sets of causes must 

be appreciated. In describing the seven communities I propose 

in the first examples to emphasize those features which make 

for orderliness, in the later to content myself with little more 

than passing reference to these and to dwell specifically upon 

departures from it; in all examples to bring out special points 

for the illustration of which particular communities are well 

suited or for which data happen to be available. For the present 

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84 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

233. Allendorf FW, Hohenlohe PA, Luikart G. 

2010. Genomics and the future of conservation genetics.

Nature Reviews Genetics 11:697-709.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: adaptation, physiological, animals, 

biodiversity, forecasting, genetic variation, population genetics, review

Abstract: We will soon have complete genome sequences 

from thousands of species, as well as from many individuals 

within species. This coming explosion of information will 

transform our understanding of the amount, distribution and 

functional significance of genetic variation in natural populations. Now is a crucial time to explore the potential implications 

of this information revolution for conservation genetics and to 

recognize limitations in applying genomic tools to conservation issues. We identify and discuss those problems for which 

genomics will be most valuable for curbing the accelerating 

worldwide loss of biodiversity. We also provide guidance on 

which genomics tools and approaches will be most appropriate 

to use for different aspects of conservation.

Link: http://www.ncbi.nlm.nih.gov/pubmed/20847747

234. Aubry C, Shoal R, Erickson VJ. 2005. Grass 

cultivars: their origins, development, and use on national forests 

and grasslands in the Pacific Northwest. Portland (OR): USDA 

Forest Service. 55 p. 

Type: Government Document

Geographic Area: Pacific Northwest USA

Compilers’ Keywords: restoration, plant materials

Abstract: Grass cultivars are a distinct subset of a species, 

often intentionally bred to behave uniformly and predictably 

when grown in an environment to which the species is adapted. 

A cultivar, also called a variety or a release, is given a unique 

trade name chosen by the breeder. These single-word names—

such as “Arlington,” “Bromar” or “Secar”—most often relate 

to the place of origin, species name, cultivar characteristics, 

or an individual involved in the process. Grass cultivars have 

been used in large quantities, often without an assessment of 

the consequences. Since grass cultivars vary in their origins, 

development history, and effects on native plant populations, 

it is important to know more than the brand name and the species name when considering a seed source for revegetation. 

Although a cultivar has been developed for particular uses 

or appears to be adapted to a wide range of conditions, the 

General (215–232)—reviews, surveys, 

summaries, theory, foundations, paradigms

Research (233–285)—propagation, 

outplanting, invasion

Strategies (286–326)—frameworks, 

approach, decision support

Resources (327–338)—native plant materials, 

guidelines, handbooks

Conservation and 

Restoration

CHAPTER 2 • CONSERVATION AND RESTORATION • RESEARCH

#. Author(s), Date. Title. Source.

Type: 

Geographic Area: 

Keywords: 

Description:

Link:

location of study

keywords written by the 

author(s) or provided by the 

compilers

ID number and

information

abstract, summary, or 

portions of the introduction 

or conclusion written by the 

author(s) or provided by the 

compilers

book, book section, conference 

paper, dissertation, government 

document, journal article, 

newsletter, presentation, report, 

thesis or web page

internet address of reference

Each reference contains the following:
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 85

perennial grass varieties might also be useful in rangeland restoration if annual precipitation is over 100 mm.

236. Benayas JMR, Newton AC, Diaz A, Bullock JM. 

2009. Enhancement of biodiversity and ecosystem services by 

ecological restoration: a meta-analysis. Science 325:1121-1124.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: assessment, reference ecosystems, 

tropical biomes, environmental degradation, restoration 

success

Abstract: Ecological restoration is widely used to reverse 

the environmental degradation caused by human activities. 

However, the effectiveness of restoration actions in increasing 

provision of both biodiversity and ecosystem services has not 

been evaluated systematically. A meta-analysis of 89 restoration assessments in a wide range of ecosystem types across the 

globe indicates that ecological restoration increased provision 

of biodiversity and ecosystem services by 44 and 25%, respectively. However, values of both remained lower in restored versus intact reference ecosystems. Increases in biodiversity and 

ecosystem service measures after restoration were positively 

correlated. Results indicate that restoration actions focused on 

enhancing biodiversity should support increased provision of 

ecosystem services, particularly in tropical terrestrial biomes.

Link: http://www.sciencemag.org/content/325/5944/1121.

short

237. Bhattarai K, Bushman BS, Johnson DA, 

Carman JG. 2011. Searls prairie clover (Dalea searlsiae) 

for rangeland revegetation: phenotypic and genetic evaluations. 

Crop Science 51:716-727.

Type: Journal 

Geographic Area: Western USA

Compilers’ Keywords: legume, rangeland restoration, genetic diversity, phenology, forb, common garden

Abstract: Few North American legumes are available for 

use in rangeland revegetation in the western USA, but Searls 

prairie clover [Dalea searlsiae (A. Gray) Barneby] is one that 

holds promise. Commercial scale seed production of this species could address the issues of unreliable seed availability and 

high seed costs associated with its wildland seed collection. To 

evaluate its utility for revegetation, we collected Searls prairie clover at 20 locations across Utah and Nevada. Amplified 

fragment length polymorphisms (AFLP) and morphological 

and phenotypic traits (measured in common-garden plots) 

were used to clarify the role of evolutionary forces responsible for its genetic structure. Collections were evaluated for 

dry-matter yield, inflorescence weight, number of inflorescences, plant height, foliage diameter, flowering date, aciddetergent fiber, neutral-detergent fiber, and crude protein at two 

material may not necessarily be suitable or optimal for all situations. Use of such material may have long-term and possibly irreversible genetic and ecological effects. Therefore it is 

essential to have a thorough understanding of plant material 

genetic origins, biological attributes, and level of compatibility 

with management objectives. In this paper we address the origins, and development of grass cultivars that have been used 

on national forests and grasslands in the Pacific Northwest. The 

genetic and ecological consequences of their use are discussed 

as well as recommendations for selection of plant materials for 

restoration projects.

Link: http://www.fs.fed.us/wildflowers/nativeplantmaterials/

documents/cultivars_maindoc_040405_appendices.pdf

235. Bailey DW, Tabini RA, Waldron BL, Libbin JD, 

Al-Khalidi K, Alqadi A, Oun MA, Jensen KB. 2010.

Potential of Kochia prostrata and perennial grasses for rangeland restoration in Jordan. Rangeland Ecology & Management 

63:707-711.

Type: Journal 

Geographic Area: Jordan, Middle East

Keywords: alternate forage sources, crested wheatgrass, forage kochia, Middle East, Russian wildrye, Siberian wheatgrass

Abstract: Six varieties of forage kochia (Kochia prostrata [L.] 

Schrad.), two Atriplex shrubs native to North America, and 

four drought tolerant perennial grass varieties were seeded and 

evaluated under arid rangeland conditions in Jordan. Varieties 

were seeded in December 2007 and evaluated in 2008 and 

2009 at two sites. Conditions were dry with Qurain receiving 

110 mm and 73 mm and Tal Rimah receiving 58 mm and 43 

mm of annual precipitation during the winters of 2007/2008 

and 2008/2009, respectively. Plants were more abundant and 

taller (P < 0.001) at Qurain than Tal Rimah in 2008. Forage 

kochia frequency was 48% and 30% in 2008 at Qurain and Tal 

Rimah, respectively. However, no seeded plants were observed 

at Tal Rimah in 2009, suggesting that 58 mm and 43 mm of annual precipitation are insufficient to allow plants to persist over 

multiple years. At the wetter site, forage kochia abundance in 

2009 was similar (P = 0.90) to that observed in 2008 and plant 

height increased (P < 0.001) from 2008 (14.4 cm ± 1.1 SE) to 

2009 (38.4 cm ± 1.1 SE). Sahro-select and Otavny-select were 

the most abundant forage kochia varieties (P = 0.05), suggesting that these experimental lines could be more adapted to the 

environmental conditions of Jordan than the commercially 

available cultivar Immigrant. Frequency of perennial grass varieties declined (P < 0.001) at Qurain from 82% ± 4 SE to 39% 

± 4 SE between 2008 and 2009, respectively. Among grasses, 

Siberian wheatgrass had better stands than crested wheatgrass, 

with Russian wildrye being intermediate. Based on this study, 

forage kochia appears to have great potential for establishing palatable perennial shrubs in arid rangeland conditions in 

Jordan if annual precipitation is at least 70 mm. Arid-adapted 

CHAPTER 2 • CONSERVATION AND RESTORATION • RESEARCH
86 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

and negatively correlated with latitude (r = 20.47, P = 0.01). 

These results are beneficial in identifying basalt milkvetch accessions that hold promise for plant improvement efforts.

239. Bockelmann AC, Wels T, Bakker JP. 2011.

Seed origin determines the range expansion of the clonal grass 

Elymus athericus. Basic and Applied Ecology 12:496-504.

Type: Journal 

Geographic Area: Europe

Keywords: invasion, seedling, herbivory, competition, habitat 

adaptation, salt marsh

Abstract: The recent invasion of clonal grasses to novel habitats poses a threat to biodiversity in various habitats. Elymus 

athericus, a clonal grass of north-western European salt marshes, is currently increasing in abundance and invading new habitats. In this study, we analyzed controlling factors for seedling 

establishment of E. athericus in frequently flooded low marsh 

habitats. Here, biotic and abiotic conditions are very different from the conditions of the parental sites with established 

populations higher up in the marsh. Hence, we hypothesized 

that seedling establishment at the expanding low marsh edge 

would depend on the parental origin (either through maternal 

effects or heritable local adaptation). We further hypothesized 

that seedling origin interacts with biotic factors such as herbivory and competition as well as with abiotic factors like inundation frequency. We tested the dependence of seedling survival, growth and vegetative reproduction on these factors in a 

factorial transplant experiment on Schiermonnikoog. Survival 

was high, with 77% of the planted seedling surviving until 

the end of the experiment. Biotic factors had a much stronger 

effect on seedling growth and mortality than parental origin 

and were independent of inundation. However, parental origin 

strongly interacted with herbivory and competition, with seedlings performing better under the conditions that resembled 

their parental site. We conclude that seedlings of E. athericus, 

a species that was previously thought to occur only in mid- to 

high marsh elevation, can establish at a frequently inundated 

low-marsh sites. Long term survival and further invasion will 

primarily depend on biotic factors in interaction with seed 

origin. Our results suggest that next to herbivory, limitation of 

seeds adapted to colonizing conditions is likely to slow down 

range expansion.

240. Bonner FT. 1990. Storage of seeds: potential and 

limitations for germplasm conservation. Forest Ecology and 

Management 35:35-43.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: seed germination, forestry, recalcitrant seeds, orthodox seeds, seed collection

common-garden locations in northern Utah. Collections from 

southern Utah and eastern Nevada exhibited high phenotypic 

values, whereas collections from western Nevada and northwestern Utah had low phenotypic values. Collections from 

northwestern Utah were genetically differentiated from those 

of southern Utah and Nevada via AFLP markers. Strong isolation by distance between collections suggests that genetic drift 

and gene flow are important factors in determining population 

structure in Searls prairie clover.

Link: http://www.fs.fed.us/rm/boise/research/shrub/

Links/2011papers/bhattaraietall2011.pdf

238. Bhattarai K, Johnson DA, Jones TA, Connors 

KJ, Gardner DR. 2008. Physiological and morphological 

characterization of basalt milkvetch (Astragalus filipes): basis 

for plant improvement. Rangeland Ecology & Management 

61:444-455.

Type: Journal 

Geographic Area: Western USA

Keywords: Astragalus filipes, restoration, revegetation, sagebrush steppe, western United States

Abstract: Astragalus filipes Torr. ex A. Gray (basalt milkvetch 

or threadstalk milkvetch) is a legume that is widely distributed 

in western North America and holds promise for revegetation 

and restoration programs in the western United States. Seed of 

67 accessions was collected in 2003 from Utah, Nevada, Idaho, 

Oregon, California, and Washington. Field-collected forage 

samples from these accessions had non-detectable or low 

levels of selenium, swainsonine, and nitrotoxins. Accessions 

were evaluated at Providence and Millville in northern Utah in 

2005 and 2006. At Providence accessions from north-central 

Oregon exhibited comparatively high biomass yield in summer and fall during both years. Basalt milkvetch accessions 

with low biomass generally had high crude protein concentration. Acid-detergent fiber and neutral-detergent fiber were 

positively correlated with biomass yield (r = 0.42, P = 0.0001; 

r = 0.57, P = 0.0001, respectively). At Millville accessions 

from north central Oregon exhibited comparatively high biomass and seed yield. Seed weight per 100 seeds varied among 

basalt milkvetch accessions in both years at Millville. Plants at 

Millville treated with imadicloprid insecticide had greater seed 

yields than nontreated plants in 2006, but not in 2005. When 

averaged across sites and years, a high correlation between 

number of stems and biomass (r = 0.82, P = 0.0001) indicated 

that number of stems is a reliable predictor of high biomass 

and seed yield. Principal component analysis of seven consolidated plant traits identified two principal components that accounted for 60% and 15% of the variation among accessions. 

The first principal component was negatively correlated with 

elevation (r = 20.71, P = 0.01) and positively correlated with 

latitude (r = 0.46, P = 0.01). The second principal component 

was positively correlated with elevation (r = 0.36, P = 0.01) 

CHAPTER 2 • CONSERVATION AND RESTORATION • RESEARCH
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 87

Potential range shifts resulting from global climate change 

pose additional threats to these high-elevation species, as suitable climates may no longer exist on the mountains where they 

grow. The combined impacts of these threats have necessitated 

active management and conservation activities. While several 

high-elevation five-needle pines have been well studied, large 

information gaps exist regarding the genetic diversity and population structure of others. This information is crucial for the 

development of conservation management strategies. In this 

report, information on genetic diversity, population structure, 

and strategies for gene conservation is presented and information gaps identified for North America’s high-elevation fiveneedle pines.

Link: http://www.treesearch.fs.fed.us/pubs/38207

242. Bowles ML, Betz RF, DeMauro MM. 1993.

Propagation of rare plants from historic seed collections: implications for species restoration and herbarium management.

Restoration Ecology 1:101-106.

Type: Journal 

Geographic Area: Central USA

Compilers’ Keywords: milkweed, milkvetch, germination, 

rare plants, Asclepias lanuginosa, Astragalus neglectus, Asclepias 

meadii, Astragalus tennesseensis, seed viability, seed storage

Abstract: Herbaria are potentially important repositories of 

living seeds that could be useful for recovery of rare plant species. To examine this capacity, we tested seed germination of 

rare milkweed (Asclepias) and milkvetch (Astragalus) species 

representing different collection dates and different herbaria. 

These groups have contrasting seed characteristics, with greater potential for longevity in the nonpermeable hard-coated 

milkvetch seeds. Twelve-year-old Asclepias lanuginosa seeds 

failed to germinate. However, we achieved 45% germination from three-year-old Asclepias meadii seeds, but germination dropped to 0% after ages of four to five years. Astragalus 

neglectus seeds germinated from 97-, 48-, and 28-year-old 

herbarium specimens, and Astragalus tennesseensis seeds germinated from a four-year-old collection. Seedlings produced 

from these experiments were incorporated into ex situ garden 

populations for recovery or restoration of rare species populations. Different herbarium pest control techniques may have 

significant bearing on the viability of seeds stored on herbarium specimens. Microwaving can cause precipitous loss of seed 

viability, while deep-freezing appears to allow some seeds to 

remain viable. Potentially live seeds of rare species should be 

stored under conditions that enhance their long-term viability.

243. Burton PJ, Burton CM. 2002. Promoting genetic 

diversity in the production of large quantities of native plant 

seed. Restoration Ecology 20:117-123.

Type: Journal 

Geographic Area: Western Canada

Abstract: Objectives of seed storage are primarily either 

short-term for forestry operations, or long-term for germplasm 

conservation. Seeds can be grouped into four classes of storage characteristics: ‘true orthodox’ seeds can be stored for long 

periods at seed moisture contents of 5-10% and sub-freezing 

temperatures; ‘sub-orthodox’ seeds can be stored under the 

same conditions, but for shorter periods due to high lipid content or thin seed coats; ‘temperate recalcitrant’ seeds cannot 

be dried at all, but can be stored for 3-5 years at near-freezing 

temperatures; and ‘tropical recalcitrant’ seeds also cannot be 

dried, and they are killed by temperatures below 10-15 °C. 

Cryogenic storage in liquid nitrogen at –196 °C holds promise, but needs more research. Some genetic damage may occur 

during seed storage, but the extent and effect of this damage 

has yet to be determined. Seed storage, by both conventional 

and cryogenic technologies, offers a relatively cheap method 

of conserving a broad range of germplasm. It will likely play 

an essential, complementary role, in germplasm conservation 

as temporary conservation until ex-situ stands are established, 

and as a safety measure against disastrous losses for limited 

numbers of seedlots. To fulfill the conservation roles, seed storage life must exceed the natural interval between germination 

and seed production for the next generation. Most true orthodox and many sub-orthodox seeds offer no problem, but neither recalcitrant seed class can meet this criterion with present 

technology. However, even short gains in viability retention 

of recalcitrant species can be useful (e.g., in transporting seed 

collections from remote areas to nurseries or laboratories).

241. Bower AD, McLane SC, Eckert A, Jorgensen 

S, Schoettle AW, Aitken SN. 2011. Conservation genetics of high elevation five-needle white pines. In: Keane RE, 

Tomback DF, Murray MP, Smith CM, editors. High Five 

Symposium. Fort Collins (CO): USDA Forest Service, Rocky 

Mountain Research Station. Proceedings RMRS-P-63. p 

98-117.

Type: Government Document

Geographic Area: Western North America

Compilers’ Keywords: Pinus, restoration, genetic diversity, 

white pine blister rust, forest management

Abstract: Conservation genetics examines the biophysical 

factors influencing genetic processes and uses that information 

to conserve and maintain the evolutionary potential of species and populations. Here we review published and unpublished literature on the conservation genetics of seven North 

American high-elevation five-needle pines. Although these 

species are widely distributed across much of western North 

America, many face considerable conservation challenges: 

they are not valued for timber, yet they have high ecological 

value; they are susceptible to the introduced disease white pine 

blister rust (caused by the fungus Cronartium ribicola) and endemic-turned-epidemic pests; and some are affected by habitat 

fragmentation and successional replacement by other species. 

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88 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

CHAPTER 2 • CONSERVATION AND RESTORATION • RESEARCH

information useful towards sustaining these tree species, including strain selection for inoculation of nursery seedlings.

Link: http://www.treesearch.fs.fed.us/pubs/38191

245. De Steven D, Sharitz RR, Singer JH, Barton 

CD. 2006. Testing a passive revegetation approach for restoring coastal plain depression wetlands. Restoration Ecology 

14:452-460.

Type: Journal 

Geographic Area: Southeastern USA 

Keywords: Carolina bays, depression wetlands, drought effects, isolated wetlands, revegetation, seed banks, wetland 

restoration

Abstract: Restoration of coastal plain depressions, a biologically significant and threatened wetland type of the southeastern 

United States, has received little systematic research. Within the 

context of an experimental project designed to evaluate several 

restoration approaches, we tested whether successful revegetation can be achieved by passive methods (recruitment from seed 

banks or seed dispersal) that allow for wetland ‘‘self-design’’ in 

response to hydrologic recovery. For 16 forested depressions 

that historically had been drained and altered, drainage ditches 

were plugged to reestablish natural ponding regimes, and the 

successional forest was harvested to open the sites and promote 

establishment of emergent wetland vegetation. We sampled 

seed bank and vegetation composition 1 year before restoration 

and monitored vegetation response for 3 years after. Following 

forest removal and ditch plugging, the restored wetlands quickly 

developed a dense cover of herbaceous plant species, of which 

roughly half were wetland species. Seed banks were a major 

source of wetland species for early revegetation. However, hydrologic recovery was slowed by a prolonged drought, which 

allowed nonwetland plant species to establish from seed banks 

and dispersal or to regrow after site harvest. Some nonwetland 

species were later suppressed by ponded conditions in the third 

year, but resprouting woody plants persisted and could alter the 

future trajectory of revegetation. Some characteristic wetland 

species were largely absent in the restored sites, indicating that 

passive methods may not fully replicate the composition of reference systems. Passive revegetation was partially successful, 

but regional droughts present inherent challenges to restoring 

depression wetlands whose hydrologic regimes are strongly 

controlled by rainfall variability.

Link: http://www.srs.fs.usda.gov/pubs/ja/ja_desteven005.pdf

246. Dey DC, Jacobs D, McNabb K, Miller G, 

Baldwin V, Foster G. 2008. Artificial regeneration of major oak (Quercus) species in the eastern United States—a review 

of the literature. Forest Science 54:77.

Type: Journal 

Geographic Area: Eastern USA

Keywords: restoration, seed production, genotype

Introduction: To address the lack of commercial quantities 

of native plant seed, we embarked on a 5-year research program to collect, propagate and screen common native grasses, 

sedges, legumes, and other forbs for use in the northern interior 

of British Columbia. Starting out with very little knowledge 

about the ecology, range, or breeding systems of candidate 

plant species, the challenge was to produce large quantities of 

seed at prices below that of wild-collected seed, while maintaining a high level of genetic diversity in this plant material. 

How could we benefit from concentrated production of single, 

identified native plant species in cultivated fields, without fully 

“domesticating” these species?

244. Cripps CL, Antibus RK. 2011. Native ectomycorrhizal fungi of limber and whitebark pine: necessary for forest 

sustainability? In: Keane RE, Tomback DF, Murray MP, Smith 

CM, editors. High Five Symposium. Fort Collins (CO): USDA 

Forest Service, Rocky Mountain Research Station. Proceedings 

RMRS-P-63. p 37-44.

Type: Government Document

Geographic Area: Western USA

Compilers’ Keywords: white pine blister rust, mountain pine 

beetle, forest sustainability, Rocky Mountains, Pinus flexilis, 

Pinus albicaulis 

Abstract: Ectomycorrhizal fungi are an important component of northern coniferous forests, including those of Pinus 

flexilis (limber pine) and P. albicaulis (whitebark pine) which 

are being decimated by white pine blister rust and mountain 

pine beetles. Ectomycorrhizal fungi are known to promote 

seedling establishment, tree health, and may play a role 

in forest sustainability. The goal of this research is to discover the native ectomycorrhizal fungi associated with these 

two pines in the Rocky Mountain region. Here we report 

32 species of ectomycorrhizal fungi associated with whitebark pine, 26 with limber pine, with an overlap of 14 species (primarily suilloids). The ectomycorrhizal fungi can be 

grouped into 1. generalists, 2. western conifer associates, 3. 

calcareous species (limber pine) and 4. specialists for fiveneedle pine or stone pines (primarily suilloids). Some of the 

Suillus species occur with stone pines globally, suggesting a 

long co-evolutionary history and important ecological roles. 

Their association with limber pines is newly reported. These 

five-needle pine specialists could confer a competitive advantage over spruce and fir when present. A preliminary 

study of the physiology of the suilloid fungi reveals intraand inter-specific variation in pH/preference/tolerance in 

vitro. Strains with limber pines from calcareous sites exhibit 

a broader pH tolerance than those found with whitebark pine 

which is restricted to high elevations. It is hoped that these 

efforts contribute to an understanding of the native ectomycorrhizal fungi with whitebark and limber pine and provide 
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 89

CHAPTER 2 • CONSERVATION AND RESTORATION • RESEARCH

Type: Journal 

Geographic Area: Intermountain West USA

Keywords: Cirsium arvense, contemporary evolution, exotic, invasion, genetic differentiation, Hesperostipa comata, 

Sporobolus airoides

Abstract: Native plant individuals often persist within communities dominated by exotics but the influence of this exposure on native populations is poorly understood. Selection for 

traits contributing to competitive ability may lead to native plant 

populations that are more tolerant of the presence of exotic invaders. In this way, long-term coexistence with an exotic may 

confer competitive advantages to remnant (experienced) native populations and be potentially beneficial to restoration. In 

past studies we have documented genetic differentiation within 

native grass populations exposed to the exotic invader Russian 

knapweed (Acroptilon repens). Here, we examine populations 

of a cool-season grass, needle-and-thread (Hesperostipa comata

[Trin. & Rupr.]) and a warm season, alkali sacaton (Sporobolus 

airoides [Torr.]) collected from Russian knapweed-invaded 

sites and adjacent noninvaded sites to assess their relative competitive ability against a novel exotic neighbor, Canada thistle 

(Cirsium arvense). Experienced S. airoides (from within A. repens invasions) appear to better tolerate (accumulate biomass, 

leaf nitrogen content, and to initiate new tillers) the presence 

of a novel competitor (C. arvense). Experienced and inexperienced H. comata genets differ in their response to the presence 

of C. arvense. Relative neighbor effects of native grasses on 

C. arvense were generally greater from experienced grasses. 

The ability to compete with novel neighbors may be driven 

by general competitive traits rather than species-specific coevolutionary trajectories. Irrespective of competitive mechanisms, the conservation of native species populations within 

weed invasions may provide an important restoration tool by 

retaining unique components of native gene pools selected by 

competitive interactions with exotics.

Link: http://weedcontrolfreaks.com/wp-content/uploads/2013/03/Fererro-Serrano-2009.pdf

249. Fiske J, Tappeiner JC. 2005. An overview of key 

silvicultural information for ponderosa pine. In: Ritchie MW, 

Maguire DA, Youngblood A, editors. Ponderosa Pine: Issues, 

Trends, and Management. Albany (CA): USDA Forest Service, 

Pacific Southwest Research Station. General Technical Report 

PSW-GTR-198. p 33-47.

Type: Government Document

Geographic Area: Western USA

Compilers’ Keywords: Pinus ponderosa, review, reforestation, timber stand development, stand growth, forest 

management

Abstract: This paper provides a selected list of classical references for the important silvicultural findings for ponderosa 

Keywords: reforestation, afforestation, planting seedlings, 

competition control, direct seeding, regeneration success, 

Quercus, oak, silviculture, animal damage

Abstract: Although natural regeneration is often the best 

method for establishing new oak (Quercus spp.) stands, there 

are increasingly more situations in which high potential for oak 

regeneration failure dictates the use of artificial regeneration 

including direct seeding and planting seedlings. Additionally, 

afforestation planting programs frequently incorporate oak 

species. Artificial regeneration of oak stands is challenging for 

numerous reasons. In this article we synthesize the current state 

of knowledge regarding growing and planting the major oak 

species in the eastern United States, point out critical research 

gaps, and provide some general rowing, planting, and stand 

tending guidelines and recommendations. Adequate site preparation, careful planting of healthy, genetically adapted seed or 

seedlings of high morphological and physiological quality, and 

subsequent control of competing vegetation and browse damage are necessary actions to assure regeneration success. Oak 

seedling survival in the early years after planting or seeding is 

a poor indicator of regeneration success. Successful regeneration may be defined as having a desired proportion of the oak 

planting stock reach dominant/codominant status in the stand. 

The costs of all activities required to produce a successful oak 

tree in the future stand should be considered in economic comparison of alternative prescriptions for oak regeneration.

Link: http://www.treesearch.fs.fed.us/pubs/14223

247. Dunne RA, Dunne CG. 2002. Potential for expanded production of native rangeland seeds in western North 

America. Native Plants Journal 3:34-37.

Type: Journal 

Geographic Area: Western North America

Keywords: seed collection, market price, seed increase, field 

production, federal agencies

Abstract: As producers of native plant seeds, we contacted 

similar businesses, asked them questions about expanding 

their cultivated production of native plant seeds, and incorporated their responses with our observations to provide a “seed 

producers’ opinion” on what is limiting native plant seed production in the western U.S. and Canada. Here, we report the 

results of this survey and discuss common problems associated 

with wildland seed production from public lands and subsequent sale to the federal government.

Link: http://www.nativeplantnetwork.org/Content/Articles/3-

1NPJ34-37.PDF

248. Ferrero-Serrano Á, Hild AL, Mealor BA. 2011.

Can invasive species enhance competitive ability and restoration potential in native grass populations? Restoration Ecology 

19:545-551.
90 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Abstract: Balancing genetic gain and diversity in seed orchard crops is of a major regeneration importance. Here we 

extend our earlier work on the development of an optimization 

protocol that maximizes crops’ genetic gain at any predefined 

diversity level, considering parental reproductive output, coancestry, and inbreeding, by incorporating variation in seed 

germination as an input. Variation in seed germination capacity substantially affected seedlots’ genetic diversity by either 

under- or overestimating their effective population size; however, genetic gains were robust and their differences were negligible. The contrasting results of gain and diversity support the 

inclusion of the germination capacity, when available, in the 

optimization protocol.

252. Glass S. 1989. The role of soil seed banks in restoration 

and management. Restoration and Management 7:24-29.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: ecological conservation, seed longevity, species composition

Introduction: The role of seed banks (the reserves of viable 

seeds in the soil) is increasingly being viewed as important 

in understanding population and community dynamics and 

in planning and interpreting a broad range of conservation 

activities including ecological preservation, management and 

restoration.

Link: http://er.uwpress.org/content/7/1/24.full.pdf

253. Johnson GR, Berrang PC. 2007. Seed shatter 

dates of antelope bitterbrush in Oregon. Rangeland Ecology & 

Management 60:99-103.

Type: Journal 

Geographic Area: Oregon, USA

Keywords: Purshia tridentata, seed collection, seed maturation

Abstract: Seed shatter dates for antelope bitterbrush (Purshia 

tridentata [Pursh] DC) were estimated from collections at 

192 sites in Oregon and surrounding states. Shatter date was 

strongly correlated to elevation (r = 0.74) and an equation that 

included elevation, latitude, longitude, and longitude squared 

explained 79% of the variation in seed shatter dates. In general, 

earlier shatter dates were associated with more southerly latitudes, easterly longitudes, and lower elevations. Examination 

of climatic data confirmed the expectation that earlier shatter 

dates were associated with warmer sites. This information can 

assist those needing to schedule seed collection activities at 

multiple locations.

254. King JN, Hunt RS. 2004. Five needle pines in 

British Columbia, Canada: past, present, future. In: Sniezko 

R, Samman S, Schlarbaum SE, Kriebel HB, editors. IUFRO 

Working Party 2.02.15. Fort Collins (CO): USDA Forest 

pine, and categorizes some of the key current literature, as well 

as some of the older, lesser known but important literature. The 

paper also provides some history of scientific developments, 

and sources of further information.

Link: http://www.treesearch.fs.fed.us/pubs/27255

250. Fu Y, Thompson D, Willms W, Mackay M. 

2005. Long-term grazing effects on genetic variability in 

mountain rough fescue. Rangeland Ecology & Management 

58:637-642.

Type: Journal 

Geographic Area: Western Canada

Keywords: native grass, Festuca campestris, cattle grazing, 

genetic diversity, AFLP

Abstract: Festuca campestris Rydb. (mountain rough fescue) 

is a dominant grass species in the montane grasslands of western Canada. Little is known about the genetic diversity of this 

plant and the effects of long-term grazing on the genetics of 

populations. The amplified fragment length polymorphism 

(AFLP) technique was applied to compare the genetic diversity of fescue plants at adjacent grazed and protected areas 

for 3 populations spread across a longitudinal range: Stavely 

in the foothills, Milroy in the Rocky Mountain trench, and 

Goose Lake on the interior plateau. Five AFLP primer pairs 

were used to screen the tiller samples of about 39 plants in each 

grazed (or ungrazed) area, and 139 polymorphic AFLP bands 

were scored for each individual sample. These scored bands 

had frequencies ranging from 0.03 to 0.98 with an average of 

0.56. About 81% of the total AFLP variation resided within the 

populations. The Goose Lake population had the lowest level 

of AFLP variation, but genetically was the most distinct. Four 

AFLP bands were possibly associated with chromosomal segments significant for grazing resistance. Comparisons of AFLP 

variation between grazing and nongrazing samples revealed 

variable and relatively small impacts of the long-term grazing 

on the genetic diversity of the grazed populations. The AFLP 

variation of grazed samples was 1.5% lower at Goose Lake, 

2.2% higher at Milroy, and not different at Stavely. If developing diverse germplasm for rangeland seeding is desired, one 

should sample across geographic space rather than combining 

materials with and without historical grazing pressure.

251. Funda T, Lstibürek M, Klápštë J, El-Kassaby 

YA. 2012. Optimization of genetic gain and diversity in 

seed orchard crops considering variation in seed germination.

Scandinavian Journal of Forest Research 27:787-793.

Type: Journal 

Geographic Area: Western Canada

Keywords: seed orchard, germination capacity, optimization, 

genetic gain, genetic diversity

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 91

only peak value increased for unstratified seed. After four days 

aging, all parameters decreased. Two days of aging enhanced 

germination capacity of unstratified seed by 15%, but stratified 

seed was still 13% higher. Douglas-fir seed should be stratified before germination, but unstratified seed can be exposed 

to 40 °C and 100% humidity for two days to augment seedling 

stock during the growing season. Ex situ Douglas-fir genetic 

resource conservation, as well as more adequate representation 

of planted genotypes across the landscape, can benefit from 

two days of aging, which would ensure slowly-germinating 

genotypes are represented in the population.

Link: http://www.researchgate.net/publication/228479582_

Effects_of_stratification_and_simulated_aging_on_germination_of_Douglas-fir_seed_from_a_clonal_seed_orchard/file/

d912f51015315cf10e.pdf

256. Kramer AT, Zorn-Arnold B, Havens K. 2010.

Assessing botanical capacity to address grand challenges in the 

United States. Chicago (IL): Chicago Botanic Garden. 64 p.

Type: Report

Geographic Area: USA

Compilers’ Keywords: workshop, report, human capacity, 

management, workforce, survey

Preface: Plants are central to the future of scientific discovery, 

human well-being, and the sustainable use and preservation of 

the nation’s resources. The botanical community in the United 

States plays a mission-critical role in researching, conserving, 

and sustainably managing our plant diversity and resources. 

Botanical expertise is required to address current and future 

grand challenges and issues, including climate change mitigation, land management and wildlife habitat restoration, understanding the provision of ecosystem services, management and 

control of invasive species, and the conservation and recovery 

of rare species. Despite the fundamental role botanical capacity plays in tackling each of these issues, this report outlines 

where botanical capacity, particularly human capacity, is lacking across all sectors (government, academic, and private). In 

the United States over the past two decades, the botanical community has experienced significant changes in the demands 

placed upon it and the resources available to it. Since the early 

1990s a series of published and anecdotal reports have outlined 

declining botanical capacity in many facets of this sector. This 

includes declines in human resources like botanical training and 

expertise, financial and management level support for research, 

education and application, and the loss of infrastructure such as 

herbaria. The nation’s science and land management agenda is 

suffering as a result. Government agencies are losing botanical 

capacity as staff botanists retire and positions are not refilled, 

either because positions are eliminated, replaced by individuals 

without equivalent botanical training, or because there is an inability to find appropriately qualified new candidates to fill them. 

Botanical education and training likewise appears to be on the 

Service, Rocky Mountain Research Station. Proceedings 

RMRS-P-32. p 12-19.

Type: Government Document

Geographic Area: Western Canada

Keywords: genetic resistance, western white pine, whitebark 

pine, limber pine

Abstract: In British Columbia (BC), Canada, we have been 

involved with white pine and blister rust since the rust’s discovery on imported infected pines through the port of Vancouver 

in 1910. Just after the rust’s introduction, the USDA Forest 

Service established monitoring plots and species trials in BC, 

but these were abandoned when the rust became well established in the USA. Resistance research began again in 1946 

with a collection of western white pine (Pinus monticola

Dougl. ex D. Don) seed that was sent to Ontario for testing. 

In about 1950 grafted plus trees were inoculated in a disease 

garden, but this work was also abandoned in 1960 when it 

was demonstrated that seedlings from such selections could 

be susceptible. Parent tree selection and seedling inoculation 

of open-pollinated families of western white pine began again 

in earnest in 1987. From this material we have the basis of a 

breeding and seed orchard program based on partial resistance 

mechanisms. An F1 generation is being produced for future 

research. Additionally, we are considering single gene resistance traits, such as major gene resistance (MGR), which can 

be pyramided onto the partial resistance of our breeding population. Efforts, particularly for conservation interests, are also 

being started for whitebark pine (P. albicaulis Engel.).

Link: http://www.fs.fed.us/rm/pubs/rmrs_p032/rmrs_

p032_012_019.pdf

255. Krakowski J, El-Kassaby YA. 2003. Effects of 

stratification and simulated aging on germination of Douglas-fir 

seed from a clonal seed orchard. Forest Genetics 10:65-70.

Type: Journal 

Geographic Area: Western Canada

Keywords: Pseudotsuga menziesii, germination parameters, 

stratification, aging

Abstract: Seed from 15 Douglas-fir (Pseudotsuga menziesii

(Mirb.) Franco) clones were germinated in a factorial design 

with two pre-treatments (unstratified and stratified) and seven simulated aging periods (0, 2, 4, 7, 10, 12 and 14 days). 

Simulated aging consisted of high temperature (40 °C) and 

humidity (100 % RH) exposure, which simulates physiological stresses and consequent deterioration in long term storage. 

Seed deteriorated as aging treatments lengthened; no germination occurred after 12 days. Germination parameters (capacity, peak value, speed, completeness) were calculated, and 

pre-treatment and aging effects evaluated using a mixed model 

analysis of variance. Germination completeness and speed 

were higher after two days of aging for stratified seed, whereas 

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CHAPTER 2 • CONSERVATION AND RESTORATION • RESEARCH

Link: http://www.bgci.org/usa/bcap

257. Krasowski MJ, Owens JN. 2000. Morphological 

and physiological attributes of root systems and seedling 

growth in three different Picea glauca reforestation stock.

Canadian Journal of Botany 30:1669-1681.

Type: Journal 

Geographic Area: Western Canada

Compilers’ Keywords: root characteristics, planting stock, 

forest, white spruce

Abstract: The relationship between certain morphological 

characteristics of white spruce (Picea glauca (Moench) Voss) 

planting stock (STK) and post-planting seedling performance 

was evaluated. Root system size at planting, its expansion, 

and its capacity to conduct water during the first post-planting 

weeks were determined. These characteristics were related 

to the performance of STK planted on two forest sites and 

measured for three growing seasons and to the performance 

of seedlings grown in large wooden boxes buried in the soil 

outdoors for one growing season (grown without competition 

from other vegetation). The compared STK were (i) polystyroblock grown, (ii) polystyroblock grown with chemical root 

pruning, and (iii) peat-board grown with mechanical root pruning. After three growing seasons on forest sites, seedlings with 

mechanically pruned roots grew more above ground than did 

seedlings from polystyroblock containers. This difference in 

seedling growth performance was even more significant for 

seedlings grown in wooden boxes. Of these, the mechanically 

pruned seedlings grew more not only above the ground but 

they also produced larger root systems by the end of the first 

growing season. This was despite the initially significantly 

smaller root systems of mechanically pruned seedlings, compared with the other two STK. Early (5–7 weeks after planting) 

post planting root expansion patterns in the three STK were 

significantly different, with the roots of mechanically pruned 

seedlings growing less than the roots in the other two STK. In 

spite of this, pressure-probe measured hydraulic conductivity 

and water flux through root systems increased during the first 

post-planting weeks in mechanically pruned seedlings while 

declining or changing little in the other two STK. It was concluded that root system size at planting and its early postplanting expansion did not relate well to the root system hydraulic 

properties or to the post-planting seedling growth performance.

258. Kujawski J, Ogle D. 2005. Not your grandpa’s 

cultivars: the new conservation releases. Native Plants Journal 

6:49-51.

Type: Journal 

Geographic Area: USA

Keywords: pre-varietal, tested, selected, source identified, 

germplasm, restoration

decline, with many botany departments at universities being 

subsumed into more general or interdisciplinary departments, 

and subsequently losing resident expertise as professors retire 

and are replaced by individuals without botanical expertise. 

Organizations in the private sector (e.g., botanic gardens and 

other non-profit conservation organizations, as well as for-profit 

businesses and self-employed individuals) are filling these widening gaps in capacity, providing botanical training, expertise, 

application, and infrastructure where it otherwise would not 

exist. Though there are ongoing concerns about funding and 

program sustainability, organizations in the private sector are 

poised to do more with additional resources and the right partnerships. Prior to this project, it was unclear exactly where the 

most critical gaps existed and which sector was most capable 

of filling them in both the short and long-term. Funding from 

the National Fish and Wildlife Foundation was awarded to the 

Chicago Botanic Garden to carry out a one year project to assess the nation’s current and future botanical capacity to conduct 

research in the plant sciences, to educate the public, train the 

next generation of plant scientists, and to conserve and manage 

the nation’s native plant species and habitat. In conducting this 

assessment, we utilized all background information available 

relating to botanical capacity (education, training, research, application, and infrastructure) in the United States. We conducted 

literature searches and obtained documents on plant science 

education, research, and application. With this information, and 

in consultation with members of an established Advisory Board 

and other individuals in the botanical community, we developed 

and conducted a series of seven on-line surveys. This included 

surveys for individuals involved in plant science research, education, or natural resource management at 1) federal government 

agencies; 2) state heritage programs; 3) other regional, state, 

county or city government agencies; 4) non-profit organizations; 

5) self-employed and for-profit companies; 6) graduate school 

(master’s and doctoral graduate students); and 7) academia 

(faculty and administrators). Surveys focused primarily but not 

exclusively on the humanly components of botanical capacity, 

and were open and publicly available for 8 weeks during the 

summer of 2009, with requests for participation sent via print 

and electronic means (e.g., the Botanical Society of America’s 

Plant Science Bulletin, Facebook, websites, email, and through 

plant science, conservation, ecology and related list serves). We 

registered more than 1,500 survey respondents representing all 

50 states, an indication that this topic is important and of interest 

throughout the United States. Survey results were an important 

source of information for this report, as the last time a survey 

was carried out that specifically targeted the botanical community in the United States was in 1989. Most surveys have been 

focused on a single sector (primarily the research/training components of the academic sector). To our knowledge, this is the 

first time multiple sectors of the botanical community (e.g., the 

entire pipeline from education and training to research, application, and employment) in the United States have been surveyed 

simultaneously.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 93

CHAPTER 2 • CONSERVATION AND RESTORATION • RESEARCH

260. Leif JW, Durling JC, Burgdorf DW. 2011.

Comparison of seed germination techniques for common elderberry (Sambucus nigra L. ssp. canadensis). Native Plants Journal 

12:132-135.

Type: Journal 

Geographic Area: USA

Keywords: American black elderberry, stratification, scarification, sulfuric acid, Caprifoliaceae

Abstract: Soaking common elderberry seeds (Sambucus nigra L. ssp. canadensis (L.) R. Bolli [Caprifoliaceae]) in sulfuric 

acid followed by a 60-d stratification, or subjecting common 

elderberry seeds to a 60-d warm, moist treatment followed by 

a 90-d stratification, significantly increased seed germination 

of accession 9084126 common elderberry seeds. Stratification 

alone was less effective in promoting germination than was 

acid scarification followed by stratification or warm, moist 

treatment followed by stratification. Germination of common 

elderberry seeds soaked in hot water was not significantly different from the control treatment.

Link: http://www.plant-materials.nrcs.usda.gov/pubs/mipmcrj10583.pdf

261. Loch DS, Boyce KG. 2003. Balancing public and 

private sector roles in an effective seed supply system. Field 

Crops Research 84:105-122.

Type: Journal 

Geographic Area: Global

Keywords: seed supply systems, public sector, private sector, 

seed quality, intellectual property

Abstract: A seed supply system can be defined in general 

terms as that combination of components, processes and their 

organisation, along with the interactions and support involved 

in the production and marketing of seeds of one or more species to a particular user–client grouping in an ongoing way. 

Seed supply systems come in a variety of forms, and operate 

at a range of levels (national, regional and local) and with different plant materials used for different purposes. In all cases, 

however, the public and the private sectors have complementary roles in ensuring that there is an effective seed supply 

system, which can meet consumer needs in terms of the range 

of plant materials available and the quantities and quality of 

seed required at an affordable price. Nationally, it is the government that sets the policy framework within which seed supply systems must operate, and also provides some supporting 

infrastructure. The private sector then is responsible for the 

timely and cost-effective production and delivery of seed to 

end users/clients within the policy framework that has been 

set by government. The balance in terms of public–private 

sector roles and responsibilities differs from place to place 

and changes over time. This paper discusses the factors that 

help shape what is an effective seed supply system under a 

Abstract: Three new types of conservation plant releases 

are now available from USDA Plant Materials Centers as 

less thoroughly tested but still useful alternatives to traditional cultivars. These “pre-varietal” releases include “tested,” “selected,” and “source-identified” plant material and 

have distinct labeling. They help meet strong customer demand for native plant material that is available more quickly 

than it takes to develop and test a cultivar for a particular 

conservation use.

Link: http://npj.uwpress.org/content/6/1/49.short

259. Lambert JK. 2008. Willamette Valley prairie restoration: the Native Seed Network and Sidalcea [Thesis]. Corvallis 

(OR): Oregon State University. 95 p.

Type: Thesis

Geographic Area: Oregon, USA

Compilers’ Keywords: applied science, conservation biology, native seed network, population genetics

Abstract: My internship was conducted with the Native Seed 

Network program within the Institute for Applied Ecology, a 

non-profit organization located in Corvallis, whose focus is 

education, applied research and active restoration and of native 

ecosystems. My internship goals were to increase my experience in applied restoration ecology and conservation biology 

with the emphasis on native plant communities in particular 

the endangered prairie communities of the Willamette Valley. 

These goals were met through my tenure with the Native 

Seed Network’s native seed collection and increase program 

during the 2007 field season. This work provided me strong 

experience with a process necessary for genetically conservative restoration and provided the NSN with a willing intern to 

complete their goals for 2007. Additional work I accomplished 

included a morphometric analysis of the native Willamette 

Valley prairie forb Sidalcea campestris which served as one 

constituent towards resolving the general ambiguity of this 

focal species for prairie restoration. Included in this analysis 

is a presentation and my interpretation of an initial molecular 

phylogenetic analysis on a selection of S. campestris accessions by Liston and deFeniks (unpublished) that may aid in 

the future resolution of S. campestris and other congeners in 

the Willamette Valley. The purpose of the morphological research was not mastery of population genetics or statistics but 

an exercise in concentration and achievement towards a more 

integrated and advanced application of taxonomic work oriented towards preserving biological diversity. This entire report 

completes the written requirement for the Professional Science 

Masters degree in Applied Systematic through the Botany and 

Plant Pathology Department at Oregon State University.

Link: http://scholarsarchive.library.oregonstate.edu/xmlui/bitstream/handle/1957/13792/Joshua.Lambert.2008.BPP-PSM.

pdf?sequence=1
94 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Keywords: cattle grazing, Inter-Simple Sequence Repeat 

markers, Festuca idahoensis, Monte Carlo procedure

Abstract: The effect of cattle grazing on the genetic structure of native grass populations has received little attention. 

We investigated the effect of cattle grazing on genetic variation in Idaho fescue (Festuca idahoensis Elmer) using ISSR 

(inter simple sequence repeat) DNA markers. The ISSR 

markers are hypervariable and are generally interpreted as 

being selectively neutral. Idaho fescue tillers were sampled 

from inside (N = 31) and outside (N = 34) a 64-year-old 

cattle exclosure in southeastern Oregon. We extracted DNA 

and used 2 ISSR primers to determine the genotypes for 

grazed and ungrazed plants at 60 variable loci. No statistically significant differences were observed between grazed 

and ungrazed samples for percent polymorphic loci (grazed 

= 85%; ungrazed = 80%), mean expected heterozygosity 

(grazed = 0.1393; ungrazed = 0.1365), or for a measure of 

loci dissimilarity (grazed = 0.506; ungrazed = 0.536). We 

also found that the ungrazed individuals sampled inside the 

exclosure were not significantly genetically differentiated 

from the grazed individuals sampled outside the exclosure 

(Gst = 0.0008 averaged across all loci). Our results differ 

from past studies that found demographic and physiological 

differences between Idaho fescue inside and outside of grazing exclosures at the same site. Our results mirror those of 

other researchers who have also failed to detect genetic differences at marker loci in response to grazing. We propose 

that either the mechanisms that must be present to cause 

changes in neutral genetic variation are not affected by cattle 

grazing for Idaho fescue at this site, or that any effects of 

grazing on neutral genetic variation were overwhelmed by 

gene flow between the grazed and ungrazed samples.

Link: https://journals.uair.arizona.edu/index.php/jrm/article/

download/12431/11710

264. Mealor BA, Hild AL. 2006. Potential selection in 

native grass populations by exotic invasion. Molecular Ecology 

15:2291-2300.

Type: Journal 

Geographic Area: USA

Keywords: AFLP, genetic differentiation, genomics, invasive 

species, native grasses, selection

Abstract: Ecological impacts of invasive plant species are 

well documented, but the genetic response of native species 

to invasive dominance has been often overlooked. Invasive 

plants can drastically alter site conditions where they reach 

dominance, potentially exerting novel selective pressures on 

persistent native plant populations. Do native plant populations 

in old exotic invasions show evidence of selection when compared to conspecific populations in adjacent, noninvaded areas? 

We employ amplified fragment length polymorphism (AFLP) 

analysis to screen a large number of loci from two native grass 

particular set of circumstances, but one that may not necessarily be as effective in other countries/regions or with a different 

product mix. It also considers the forces that lead to a gradual 

evolution of the system over time and to recent and more radical changes in major developed countries. The latter shift has 

been brought about by the commercial opportunities presented 

by genetic engineering and legal protection of the intellectual 

property (IP) in new plant varieties, coupled with a shift in 

government policy towards deregulation of the economy (and 

so greater industry self-regulation in areas like quality assurance). These factors have led to far-reaching structural changes 

in relation to commercial seed production and delivery in the 

developed world over the past decade. The recent changes in 

seed supply arrangements were focused initially on high volume arable food crops in the larger developed economies of 

North America and western Europe. However, they are now 

gradually moving through to lower volume seed markets and 

to smaller, less developed economies where they will have different implications for the development of effective country 

specific seed supply policies.

Link: http://startinternational.org/library/archive/files/publicprivate-seed-sector_d371f58830.pdf

262. Losey JE, Vaughan M. 2006. The economic value of 

ecological services provided by insects. BioScience 56:311-323.

Type: Journal 

Geographic Area: USA

Keywords: ecological services, economic value, conservation, biodiversity, environmental policy

Abstract: In this article we focus on the vital ecological services provided by insects. We restrict our focus to services 

provided by “wild” insects; we do not include services from 

domesticated or mass-reared insect species. The four insect 

services for which we provide value estimates—dung burial, 

pest control, pollination, and wildlife nutrition—were chosen 

not because of their importance but because of the availability 

of data and an algorithm for their estimation. We base our estimations of the value of each service on projections of losses 

that would accrue if insects were not functioning at their current level. We estimate the annual value of these ecological 

services provided in the United States to be at least $57 billion, 

an amount that justifies greater investment in the conservation 

of these services.

Link: http://xerces.org/wp-content/uploads/2008/09/economic_value_insects.pdf

263. Matlaga D, Karoly K. 2004. Long-term grazing 

effects on genetic variation in Idaho fescue. Rangeland Ecology 

& Management 57:275-279.

Type: Journal 

Geographic Area: Oregon, USA

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 95

266. Merritt DJ, Dixon KW. 2011. Restoration seed 

banks—a matter of scale. Science 332:424-425.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: conservation, long-term management, provenances, seed collection

Compilers’ Summary: Comment on the current state of restoration in relation to seed banks and scale. The authors preface the article with the fact that seed banks do not have enough 

seed for large-scale restoration projects. Seed banks must shift 

from being “stamp-collections” of species to collections that can 

provide tons of seeds and the expertise to improve restoration 

efforts. Further, in the seed bank system, seed source, availability, or management and use of seeds are rarely considered 

in collections. The lack of information about each seed lot can 

lead to storage and planting failures. Some needs are seed farming of native species, seed technology for effective use and site 

delivery, genetic analysis tools to address provenances in seed 

farming, refinement of seed collection and utilization strategies, 

and transfer of knowledge on collecting, using and storing seeds. 

Link: http://www.esalq.usp.br/lcb/lerf/divulgacao/recomendados/artigos/merritt2011.pdf

267. Miller RS. 1967. Pattern and process in competition. 

Advances in Ecological Research 4:1-74.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: plant populations, spatial distribution, coexistence, species diversity

Summary: This chapter discusses the development of competition theory, which has traditionally included three stages: 

inferences drawn from observation of natural populations, 

construction of mathematical models, and laboratory experiments designed to test elements of competitive interactions 

in controlled environments. Interference to any activity either 

directly or indirectly limits a competitor’s access to a necessary resource or requirement. Different types of evidence are 

assigned for competition in nature. Principal categories are 

mutually exclusive spatial distributions without supporting 

evidence of a competitive interaction, observed or inferred 

ecological displacement in sympatric populations, and induced 

changes in distribution pattern. A considerable amount of ambiguity exists in interpretations of the results of interspecies 

competition. It is necessary to distinguish between maximum 

exploitation of the available resources by one species, and 

equitable utilization of the resources and possible coexistence 

in a mixed species system. The chapter suggests two major 

sources of species diversity. When competition is primarily 

through exploitation and the system is under strong environmental control, it is likely that fluctuations in factors affecting 

reproduction and survival will continually alter the outcome 

species (Hesperostipa comata (Trin. & Rupr.) Barkworth and 

Sporobolus airoides Torr.) that occur in old infestations of the 

invasive forb Acroptilon repens. We then compare observed 

locus by locus FST values with distributions of FST estimated 

from simulation models under expectation of neutrality. We 

also compare the proportion of loci possibly linked to selection and those not linked to selection which exhibit parallel 

trends in divergence between two community types (invaded, 

noninvaded). Few loci (H. comata, 2.6%; S. airoides, 8.7%) in 

the two native grasses may be linked to genes under the influence of selection. Also, loci linked to selection showed a 

greater portion of parallel trends in divergence than neutral 

loci. Genetic similarities between community types were less 

than genetic similarity within community types suggesting differentiation in response to community alteration. These results 

indicate that a small portion of scored AFLP loci may be linked 

to genes undergoing selection tied to community dominance 

by an invasive species. We propose that native plants in communities dominated by exotic invasives may be undergoing 

natural selection.

Link: http://www.ncbi.nlm.nih.gov/pubmed/16780441

265. Mealor BA, Hild AL, Shaw NL. 2004. Native 

plant community composition and genetic diversity associated with long-term weed invasions. Western North American 

Naturalist 64:503-513.

Type: Journal 

Geographic Area: USA

Keywords: native grasses, invasive plants, diversity, ISSR, 

Acroptilon repens, Cardaria draba, Achnatherum hymenoides, 

Hesperostipa comata, Sporobolus airoides, Poa secunda

Abstract: Many studies have assessed genetic changes in 

exotic plant species in their native and introduced ranges, 

but none have focused on genetic variation in native plant 

species in response to exotic invasion. We examine characteristics of native plant communities within and outside old 

(>25 year) invasions of Acroptilon repens (Russian knapweed) 

and Cardaria draba (hoary cress). We also document genetic 

variability of 4 native grass populations (Hesperostipa comata

[needle and thread], Achnatherum hymenoides [Indian ricegrass], Sporobolus airoides [alkali sacaton], and Poa secunda

[Sandberg bluegrass]) from 2 areas: adjacent to and within 

weed invasions. Native plant species richness and diversity 

did not differ between invaded and noninvaded areas. Intersimple sequence repeat (ISSR) analysis of individual native 

perennial grasses of each of the 4 species suggests that populations exposed to long-term coexistence with exotics may 

differ from adjacent noninvaded populations. We suggest that 

future research efforts should focus on intraspecific diversity 

of native plant species to identify possible candidates for restoration following weed control.

Link: http://www.treesearch.fs.fed.us/pubs/40785

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CHAPTER 2 • CONSERVATION AND RESTORATION • RESEARCH

Geographic Area: Texas, USA

Keywords: herbage nutritive value, yield, wildlife, revegetation, rangeland

Abstract: Native seed mixes for rangeland seeding, prairie 

restoration, or cultivated pasture can benefit from a greater variety of forbs that more closely reflect the original vegetation 

of the southern Great Plains. Fifteen native, perennial herbaceous legumes were collected in central Texas and evaluated 

for herbage production, mineral content, and fiber concentration of established plants in research plots over 2 years. Downy 

milk-pea (Galactia volubilis [L.] Britton) was productive, regardless of rainfall, whereas prairie acacia (Acacia angustissima [Mill.] Kuntze var. hirta [Nutt.] B.L. Rob.) and Illinois 

bundle-flower (Desmanthus illinoensis [Michx.] MacMill. Ex. 

B.L. Rob. & Fernald) out-yielded others in year 3 when rainfall was the greatest. Herbage crude protein averaged approximately 100 g/kg for bush-clovers (Lespedeza spp.) compared to 

bundle-flowers (Desmanthus spp.), which exceeded 200 g/kg; 

the latter also was high in herbage phosphorus. Herbage neutral 

detergent fiber ranged from 300 to more than 500 g/kg, acid detergent fiber ranged from 140 to 360 g/kg, and acid detergent 

lignin ranged from 36 to 140 g/kg, a wide range from which 

to select if animal nutrition is a primary criterion. Seed production was evaluated within a subset of 8 entries submitted 

to periodic herbage removal or left intact throughout the season. Three bundle-flowers yielded the greatest mass and seed 

number, but were negatively affected by harvest, unlike prairie acacia. Herbage and seed characteristics indicate there are 

promising perennial herbaceous legumes in the southern Great 

Plains that can be included in native seed mixes.

271. Page MJ, Baxter GS, Lisle AT. 2006. Evaluating 

the adequacy of sampling germinable soil seed banks in semiarid systems. Journal of Arid Environments 64:323-341.

Type: Journal 

Geographic Area: Australia

Keywords: soil seed banks, species richness estimates, sample 

size, sampling strategy, seed distribution, semi-arid, first-order 

jackknife

Abstract: Quantification of soil seed is an important tool for 

understanding vegetation dynamics. However, determining 

adequate sample sizes and sampling regimes is problematic. 

The literature reports highly variable sample sizes and regimes without any generally applicable standards appearing to 

emerge. This problem is exaggerated by the heterogeneous and 

patchy nature of arid and semi-arid environments. Species accumulation curves are commonly used in floristic sampling to 

plan sampling effort and to evaluate its adequacy. However, the 

precision of the sample cannot be quantified and it is difficult 

to determine the number of species missed. We suggest an alternate approach in which a species richness estimator is compared to the actual number of species found. When applied to 

of the competitive interaction, allowing coexistence of mixed 

species populations

Link: http://www.sciencedirect.com/science/article/pii/

S0065250408603190

268. Monsen SB, Shaw NL. 2001. Development and 

use of plant resources for western wildlands. In: McArthur 

ED, Fairbanks DJ, editors. Shrubland Ecosystem Genetics 

and Biodiversity. Fort Collins (CO): USDA Forest Service, 

Rocky Mountain Research Station. Proceedings RMRS-P-21. 

p 47-61.

Type: Government Document

Geographic Area: Western USA

Compilers’ Keywords: native plant materials, biodiversity, 

restoration, native seed, review

Abstract: Concern for declines in big game habitat throughout 

the West and the pioneering work of revegetation researchers in 

the mid twentieth century led to increased use of native shrubs, 

grasses, and forbs for revegetation, and the 1975 establishment 

of the USDA Forest Service, Shrub Sciences Laboratory in 

Provo, Utah. During this period revegetation objectives shifted 

from an emphasis on production of commodities to conservation of biodiversity and ecosystem functions. Plant resource 

development altered from an agronomic approach focusing on 

plant improvement to one that incorporates ecological, genetic, 

and practical considerations. Although many problems remain, 

research, technological advances, efforts to stabilize the native 

seed industry, and improved seed testing and certification procedures are increasing our options for revegetating disturbed 

lands.

Link: http://www.treesearch.fs.fed.us/pubs/40780

269. Mortlock W. 2000. Local seed for revegetation: 

where will all that seed come from? Ecological Management 

and Restoration 1:93-101.

Type: Journal 

Geographic Area: Australia

Compilers’ Keywords: seed collection, large-scale demand, 

local adaptation

Abstract: Broad-scale revegetation requires large quantities 

of locally indigenous seed, but there are worrying signs that 

current demand for local seed may already be outstripping 

supply in some areas of Australia. Could integrated solutions 

include better regional cooperation for seed collection as well 

as the creation of seed production areas?

270. Muir JP, Taylor J, Interrante SM. 2005. Herbage 

and seed from Texan native perennial herbaceous legumes. 

Rangeland Ecology & Management 58:643-651.

Type: Journal 
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 97

CHAPTER 2 • CONSERVATION AND RESTORATION • RESEARCH

Keywords: annual cereal grains, Burned Area Emergency 

Response, native plant species

Abstract: Broadcast seeding is one of the most commonly 

used post fire rehabilitation treatments to establish ground 

cover for erosion control and mitigation of non-native plant 

species invasions. Little quantitative information is available 

on overall trends of post-fire seeding expenditures and seed 

mixes used over time in forested ecosystems in the western 

USA. We reviewed scientific articles, government publications 

and unpublished documents as well as USDA Forest Service 

Burned Area Reports to determine trends in post-fire seeding in 

forested ecosystems over time. Of 1164 USDA Forest Service 

Burned Area Reports, 380 contained information on seeding 

treatments conducted in forested ecosystems. A review of 40 

papers and 67 Burned Area Reports reporting species seeded 

between 1970 and 2007 revealed a trend of increasing use of 

native species, annual cereal grains and sterile-grass hybrids, 

with natives dominating seed mixes. According to 380 Burned 

Area Reports with data on costs and area seeded, total post-fire 

seeding expenditures have increased substantially, averaging 

U.S. $3.3 million per year spent on post-fire emergency seeding treatments in forested ecosystems that involved the Forest 

Service during the period 2000 to 2007—an increase of 192% 

compared with the average spent during the previous 30 years. 

The percentage of the total burned area seeded averaged 21% 

in the 1970s, compared with only 4% between 2000 and 2007.

Link: http://www.fs.fed.us/rm/pubs_other/rmrs_2011_peppin_d001.pdf

274. Pitman WD. 2009. Establishment and survival of native legumes on upland sites in Louisiana. Native Plants Journal 

10:241-250.

Type: Journal 

Geographic Area: Louisiana, USA

Keywords: renovation, Centrosem virginianum, Mimosa strigillosa, Neptunia lutea, Rhynchosia latifolia, Strophostyles umbellata, Tephrosia virginiana

Abstract: Field evaluations of perennial native legumes 

(Fabaceae) from Louisiana pineland ecosystems revealed differing colonization abilities among species under natural recolonization of disturbed sites. Initial establishment and growth 

of transplanted seedlings of 5 different species were good in 

an early field evaluation. In a subsequent evaluation initiated 

with scarified seeds, poor establishment of herbaceous mimosa 

(Mimosa strigillosa Torr. & A. Gray) indicated that populations 

of this species may be limited by lack of competitiveness of 

emerging seedlings. Virginia tephrosia (Tephrosia virginiana

(L.) Pers.) produced vigorous upright seedling growth with 

population expansion in only one year of these evaluations. 

Initial establishment of prairie snoutbean (Rhynchosia latifolia

Nutt. ex Torr. & A. Gray) was limited by inadequate rainfall 

in some instances, but under favorable conditions this species 

data collected in a semi-arid region of South-West Queensland, 

this method gave a good indication of sampling adequacy at 

large sample sizes. We suggest that this post hoc comparison 

of the predicted species richness (with coefficient of variation) 

to the actual number of species found would be useful in all 

studies reporting seed bank composition. This would allow the 

reader to evaluate the adequacy of the sampling effort, when 

judged against the research aims. We also investigated the deployment of sampling effort by examining structured subsets 

of our data. The results suggest that sampling effort may be 

optimized by taking fewer samples per transect in any one site, 

but selecting more transects and sites.

272. Peppin DL, Fulé PZ, Lynn JC, Mottek-Lucas 

AL, Hull Sieg C. 2010. Market perceptions and opportunities for native plant production on the southern Colorado plateau. Restoration Ecology 18:113-124.

Type: Journal 

Geographic Area: Southwestern USA

Keywords: local genotype, market perception differences, native plant market, native plant policy, web-based survey

Abstract: Increases in revegetation activities have created a 

large demand for locally adapted native plant materials (NPM) 

in the southwestern United States. Currently, there is a minimal 

supply of local genotypes to meet this demand. We investigated the potential for the initiation of a native plant market 

in the southern Colorado Plateau. Through a literature search, 

interviews, and site visits, we identified existing native plant 

markets outside of the region as useful models to help initiate 

a regional market. We used web based surveys to identify and 

analyze current and future NPM needs and concerns. Survey 

results indicate that management policy strongly drives decisions regarding the use and purchase of NPM. From a demand 

perspective, lack of availability and cost of NPM has kept purchasing minimal, despite policy changes favoring the use of 

natives. For suppliers, further development of NPM is limited 

by inconsistent and unreliable demand and lack of production 

knowledge. The knowledge and tools necessary to initiate an 

NPM market are available, but inadequate funding sources 

and insufficient information sharing hinder its development. 

Communication among producers, land managers, buyers, and 

researchers, as well as partnerships with local growers, appear 

to be vital to initiating a functional market.

Link: http://www.fs.fed.us/rm/pubs_other/rmrs_2010_peppin_d001.pdf

273. Peppin DL, Fulé PZ, Hull Sieg C, Beyers JL, 

Hunter ME, Robichaude PR. 2011. Recent trends in 

post-wildfire seeding in western U.S. forests: costs and seed mixes. International Journal of Wildland Fire 20:702-708.

Type: Journal 

Geographic Area: Western USA
98 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

length (SRL) at high (and low) temperatures, respectively. 

Although P. spicata’s greater productivity may initially make 

for better seedling establishment than E. wawawaiensis, it 

may also prove disadvantageous in competitive or highly resource limited environments where high SLA or SRL could 

be an advantage.

Link: http://www.researchgate.net/publication/231186284_

Immature_Seedling_Growth_of_Two_North_American_Native_Perennial_Bunchgrasses_and_the_Invasive_Grass_Bromus_tectorum/file/d912f5065c1749dd03.pdf

276. Rowe CLJ, Leger EA. 2011. Competitive seedlings and inherited traits: a test of rapid evolution of Elymus 

multisetus (big squirreltail) in response to cheatgrass invasion.

Evolutionary Applications 4:485-498.

Type: Journal 

Geographic Area: California, Nevada, USA

Keywords: adaptation, competition, contemporary evolution, ecological genetics, invasive species, natural selection, 

restoration

Abstract: Widespread invasion by Bromus tectorum (cheatgrass) in the Intermountain West has drastically altered native 

plant communities. We investigated whether (big squirreltail) 

is evolving in response to invasion and what traits contribute 

to increased performance. Seedlings from invaded areas exhibited significantly greater tolerance to B. tectorum competition and a greater ability to suppress B. tectorum biomass than 

seedlings from adjacent uninvaded areas. To identify potentially adaptive traits, we examined which phenological and 

phenotypic traits were correlated with seedling performance 

within the uninvaded area, determined their genetic variation 

by measuring sibling resemblance, and asked whether trait distribution had shifted in invaded areas. Increased tolerance to 

competition was correlated with early seedling root to shoot 

ratio, root fork number, and fine root length. Root forks differed among families, but none of these traits differed significantly across invasion status. Additionally, we surveyed more 

broadly for traits that varied between invaded and uninvaded 

areas. Elymus multisetus plants collected from invaded areas 

were smaller, allocated more biomass to roots, and produced 

a higher percentage of fine roots than plants from uninvaded 

areas. The ability of native populations to evolve in response to 

invasion has significant implications for the management and 

restoration of B. tectorum-invaded communities.

Link: http://naes.agnt.unr.edu/PMS/Pubs/548_2011_03.pdf

277. Samnegård U, Persson AS, Smith HG. 2011.

Gardens benefit bees and enhance pollination in intensively 

managed farmland. Biological Conservation 144:2602-2606.

Type: Journal 

Geographic Area: Sweden, Europe

was superior in seed production and population expansion. 

Individual plants of these species were rather short-lived with 

long-term population survival dependent on seed production 

and seedling recruitment. Seed germination of these legumes 

was limited by hard seedcoats. Physical scarification readily 

overcame this limitation for all species evaluated. Rhynchosia 

latifolia appears to have potential value for use in reclamation and revegetation plantings because of potential for seed 

production and establishment of persistent populations under 

favorable conditions. Mimosa strigillosa has potential to provide superior ground cover and sustainable populations, but 

limited establishment from seeds indicates that selection from 

expanded germplasm collections for seedling vigor may be 

particularly important for successful use of this species.

275. Ray-Mukherjee J, Jones TA, Adler PB, Monaco 

TA. 2011. Immature seedling growth of two North American 

native perennial bunchgrasses and the invasive grass Bromus tectorum. Rangeland Ecology & Management 64:358-365.

Type: Journal 

Geographic Area: Intermountain West USA

Keywords: absolute growth rate, bluebunch wheatgrass, relative growth rate, Snake River wheatgrass, specific leaf area, 

specific root length

Abstract: Pseudoroegneria spicata (Pursh) A. Love and 

Elymus wawawaiensis J. Carlson & Barkworth are two native perennial grasses widely used for restoration in the 

Intermountain West. However, the rapid establishment and 

spread of Bromus tectorum L., an invasive annual grass, has 

led to a decline in the abundance of native perennial grasses. 

Proliferation of B. tectorum has been attributed to its early 

germination, superior cold-temperature growth, profuse root 

production, and high specific leaf area (SLA). To enhance 

restoration success, we compared B. tectorum to commercially available plant materials of two perennial rangeland 

bunchgrasses, P. spicata (cv. Whitmar, cv. Goldar, and 

Anatone Germplasm) and E. wawawaiensis (cv. Secar), for 

germination, seedling morphological traits, and growth rates 

at the immature seedling stage. We monitored germination 

and immature seedling growth in a growth chamber in two 

separate experiments, one under low (5/10 °C) and the other 

under high (15/20 °C) day/night temperatures. Compared 

to the average of the two perennials, B. tectorum was 93% 

(77%) greater at high (and low) temperature for root:shoot 

length ratio, but only 14% (14%) greater for root:shoot biomass ratio and 12% (19%) lower for SLA. This suggests that 

B. tectorum’s substantial investment in surface area of roots, 

rather than in shoot length, root biomass, or leaf area, may 

be responsible for the annual’s success at the early seedling 

stage. Compared to E. wawawaiensis, P. spicata averaged 

65% (41%) higher shoot biomass, 39% (88%) higher root 

biomass, and 70% (10%) higher absolute growth rate, but 

25% (15%) lower SLA and 15% (36%) lower specific root 

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279. Smith FS, Lloyd-Reilley J, Ocumpaugh WR. 

2010. South Texas Natives: a collaborative regional effort to 

meet restoration needs in south Texas. Native Plants Journal 

11:252-268.

Type: Journal 

Geographic Area: Texas, USA

Keywords: USDA NRCS Plant Materials Program, restoration, revegetation, Rio Grande Plains, Sand Plains, Gulf 

Prairies and Marshes

Abstract: South Texas Natives (STN) is an initiative started 

in 2001 to develop and promote native plants for the restoration and reclamation of public and private lands in south Texas. 

At the urging of concerned conservationists and private landowners, STN has developed commercially viable sources of 

native seed and conducted research to discover effective restoration strategies that can be used by private landowners and 

government agencies to restore native plant communities. STN 

grew out of a strong partnership among the USDA Natural 

Resources Conservation Service E “Kika” de la Garza Plant 

Materials Center, Texas AgriLife Research, Rio Farms Inc, 

Caesar Kleberg Wildlife Research Institute, and private landowners of south Texas. Plant development efforts center on the 

release of commercially viable, multiple origin germplasms 

selected with an awareness of the commercial requirements 

for production but grounded in genetic and ecosystem function 

parameters. In addition to plant development efforts, extensive 

restoration and revegetation research is conducted by STN to 

develop usable methodology and much-needed guidelines for 

restoration in south Texas. Current restoration research emphasis centers on providing techniques for the diversification of 

areas dominated by invasive exotic grasses.

280. Sniezko R, Kegley A, Danchok R, Long S. 

2007. Variation in resistance to white pine blister rust among 

43 whitebark pine families from Oregon and Washington - 

Early results and implications for conservation. In: Goheen 

EM, Sniezko R, editors. Whitebark Pine: A Pacific Coast 

Perspective. Portland (OR): USDA Forest Service, Pacific 

Northwest Region. R6-NR-FHP-2007-01. p 82-97.

Type: Government Document

Geographic Area: Pacific Northwest USA

Compilers’ Keywords: resistance, seed collection, inoculation, pest management, Cronartium ribicola

Abstract: All nine North American species of white pines are 

susceptible to the introduced, invasive pathogen Cronartium 

ribicola, the cause of white pine blister rust. Whitebark pine 

is considered one of the most susceptible species. Genetic resistance is considered a cornerstone for survival to this pathogen. Fortunately, all of the native species of white pines have 

some level of resistance. Evaluation of resistance in Oregon 

and Washington families of whitebark pine has only recently 

Keywords: Apoidea, Bombus, Campanula, pollinator, seed 

set, agriculture, landscape

Abstract: The recent loss of pollinating insects and outcrossing plants in agricultural landscapes has raised concern 

for the maintenance of ecosystem services. Wild bees have 

been shown to benefit from garden habitats in urban and 

suburban areas. We investigated the effects of distance from 

garden habitats on wild bees and seed set of a native out 

crossing plant Campanula persicifolia, in intensively managed agricultural landscapes in southern Sweden. Bee abundance and species richness, as well as plant seed set, were 

higher closer to gardens (<15 m) than further away (>140 

m). This highlights private gardens as a landscape wide resource for pollinators but also the lack of sufficient pollination of wild plants in contemporary agricultural landscapes.

278. Shaukat SS, Siddiqui IA. 2004. Spatial pattern 

analysis of seeds of an arable soil seed bank and its relationship 

with above-ground vegetation in an arid region. Journal of Arid 

Environments 57:311-327.

Type: Journal 

Geographic Area: Pakistan, Southern Asia

Keywords: spatial pattern, seed bank, weeds, vegetation, 

dispersal

Abstract: The composition and spatial pattern of weeds 

was analysed in an arable-soil seed bank after the harvest of 

wheat. Four hundred cores were collected from a 10x10m 

grid in April 1999. Three indices of pattern detection (variance/mean ratio, Lloyd’s index of mean crowding, and 

Morisita’s index of aggregation) and Moran’s I statistic of 

spatial autocorrelation were computed. The above-ground 

vegetation of the corresponding area was sampled by 100 

randomly placed quadrats in March and September 1999 

and variance/mean ratio of all the species was calculated. 

The seed bank of the field was dominated by annuals including Convovulus arvensis, Amaranthus viridis, Anagallis 

arvensis, Chenopodium album and Conyza canadensis. In all 

27 species were recorded from the seed bank. For most of 

the species with a mean higher than 0.05 per core, the seed 

pattern (with two exceptions) was aggregated as indicated 

by various indices of pattern detection. Among the common 

species, only Amaranthus viridis exhibited a random pattern. Aggregated pattern of seeds was generally due to seeds 

settling in the neighbourhood of the mother plant and the 

clumped distribution of plant species in the overlying vegetation. Suggestions have been made regarding the improvement of the precision of the soil seed bank sample, in view 

of the aggregated distribution of seeds in soil. Seed bank 

showed considerable qualitative similarity with the aboveground vegetation, which is presumably the consequence of 

recurrent disturbances in the arable field.

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was countered by the development of a modest forest floor 

seed bank (6% of the original canopy seed bank) from burial 

of cones; this seed bank may be ecologically important if 

a fire or anthropogenic disturbance exposes these cones. If 

adequate levels of regeneration are to occur, disturbances to 

create seedbeds must occur shortly after tree mortality, before the seed banks are lost. Our findings also suggest that 

the sustained seed rain (over at least nine years) after MPB 

outbreak may be beneficial for population growth of groundforaging vertebrates. Our study adds insight to the seed ecology of serotinous pines under a potentially continental-wide 

insect outbreak, threatening vast forests adapted to regeneration after fire.

Link: http://digitalcommons.usu.edu/barkbeetles/232/

282. Thompson PA. 1976. Factors involved in the selection of plant resources for conservation as seed in gene banks. 

Biological Conservation 10:159-167.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: seed bank, seed conservation, plant 

conservation genetics, cold storage, review

Abstract: The preservation of seed in cold stores provides 

a convenient and effective method of conserving many species of plants. The ease with which collections of some species may be kept in this way poses the danger that it may be 

applied uncritically to populations or taxa to which it is less 

suited. The paper briefly reviews the major factors which 

contribute to the success of collections for a seed bank and 

derives criteria from them which may be used as a means of 

evaluating proposals for seed conservation by this means. 

The method can also be used to identify possible problems 

or to estimate relative expenditures of time and other resources which would be involved in making collections or 

looking after banked seed.

283. Tomback DF, Achuff P, Schoettle AW, 

Schwandt JW, Mastrogiuseppe RJ. 2011. The magnificent high-elevation five-needle white pines: ecological roles 

and future outlook. In: Keane RE, Tomback DF, Murray 

MP, Smith CM, editors. The Future of High-Elevation, FiveNeedle White Pines in Western North America: Proceedings 

of the High Five Symposium. Fort Collins (CO): USDA Forest 

Service, Rocky Mountain Research Station. Proceedings 

RMRS-P-63. p 2-28.

Type: Government Document

Geographic Area: Western USA

Compilers’ Keywords: mountain pine beetle, white pine 

blister rust, review, threats

Abstract: The High Five symposium is devoted to exchanging 

information about a small group of pines with little commercial 

begun; currently over 150 seedlots collected from individual 

parent trees are in resistance testing. This report summarizes 

differences in responses among 43 seedling families and one 

bulked seedlot through two years after artificial inoculation 

with blister rust. Initial infection after inoculation of threeyear-old seedlings was very high in the first set of trials; 100% 

of the seedlings developed needle lesions in the two trials reported in this paper. There were large differences among families in several traits, including percentage of trees with stem 

symptoms and survival two years after inoculation. The level 

of resistance present in some families and the frequency of resistance among the 43 families reported here is encouraging. 

A possible geographic trend in resistance is also noted. It is 

recommended that at least a subset of families be field planted 

to validate resistance ratings from this short-term screening. 

The collection and use of seed from putative resistance parent 

trees identified through this testing would be a good starting 

point for restoration efforts.

Link: http://www.fs.fed.us/r6/dorena/documents/publications/pub308.pdf

281. Teste FP, Lieffers VJ, Landhausser SM. 2011.

Seed release in serotinous lodgepole pine forests after mountain 

pine beetle outbreak. Ecological Applications 21:150-162.

Type: Journal 

Geographic Area: Western Canada

Keywords: biotic disturbance, cone burial, cone opening, 

Dendroctonus ponderosae, ground-foraging vertebrates, mountain pine beetle, natural regeneration, Pinus contorta var. latifolia, Rocky Mountain lodgepole pine, seed banks, serotiny 

(canopy seed storage), Tamiasciurus hudsonicus

Abstract: There are concerns that large-scale stand mortality due to mountain pine beetle (MPB) could greatly 

reduce natural regeneration of serotinous Rocky Mountain 

(RM) lodgepole pine (Pinus contorta var. latifolia) because 

the closed cones are held in place without the fire cue for 

cone opening. We selected 20 stands (five stands each of 

live [control], 3 years since MPB [3-yr-MPB], 6 years since 

MPB [6-yr MPB], and 9 years since MPB [9-yr-MPB] mortality) in north central British Columbia, Canada. The goal 

was to determine partial loss of serotiny due to fall of crownstored cones via breakage of branches and in situ opening 

of canopy cones throughout the 2008 and 2009 growing 

seasons. We also quantified seed release by the opening of 

forest-floor cones, loss of seed from rodent predation, and 

cone burial. Trees killed by MPB three years earlier dropped; 

3.5 times more cones via branch breakage compared to live 

stands. After six years, MPB-killed stands had released 45% 

of their canopy seed bank through cone opening, cone fall 

due to breakage, and squirrel predation. Further losses of 

canopy seed banks are expected with time since we found 

9-yr-MPB stands had 38% more open canopy cones. This 
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 101

CHAPTER 2 • CONSERVATION AND RESTORATION • RESEARCH

284. Vance NC. 2010. Evaluation of native plant seeds 

and seeding in the East-Side Central Cascades Ponderosa Pine 

Zone. Portland (OR): USDA Forest Service, Pacific Northwest 

Research Station. General Technical Report PNW-GTR-823. 

85 p.

Type: Government Document

Geographic Area: Pacific Northwest USA

Keywords: postfire rehabilitation, native forb, native grass, 

Pinus ponderosa/Purshia tridentate/Festuca idahoensis plant association, seeds, seedlings east-side central Oregon Cascades

Abstract: In dry, open coniferous forests of the montane West, 

stand-replacing wildfires and land use activities alter the composition and abundance of native grasses and forbs by degrading the habitat and accelerating the invasion of exotic annuals. 

On these lands, native forbs and grasses delayed or prevented 

from recovery by natural processes may require intervention 

through supplementary seeding. However, effective seeding of 

native plants requires that their seed traits and the potential outcome of the seeding be better understood. This study evaluated 

seeds and seedlings of 13 native forbs and 5 grasses common 

in the dry Pinus ponderosa/Purshia tridentata/Festuca idahoensis plant communities east of the Oregon Cascades crest and 

their potential for establishment in a landscape altered by past 

grazing and a recent, stand-replacing wildfire. Their potential 

for germination and establishment was examined in the laboratory and in 20 test plots located within the burned boundary of 

a 2002 wildfire. Seed collection, handling, testing, and sowing 

procedures are described. Data on seedling emergence, height, 

second-year survival and cover are presented in tables and figures. Onsite emergence and early growth data helped to reveal 

cause of early mortality related to biological and site conditions, including invasive competition, and differences in early 

growth and site occupancy. The study supports the efficacy of 

using multiple and functionally diverse species in a seeding 

program. Additional information on each of the 18 species is 

included in an appendix.

Link: http://www.treesearch.fs.fed.us/pubs/36921

285. Walters C, Wheeler LM, Grotenhuis JM. 

2005. Longevity of seeds stored in a genebank: species characteristics. Seed Science Research 15:1-20.

Type: Journal 

Geographic Area: USA, Global

Keywords: storage, viability, longevity, conservation, genebank, Avrami, composition, evolution, genetic diversity, seed

Abstract: Seeds of different species are believed to have characteristic shelf lives, although data confirming this are scarce, 

and a mechanistic understanding of why this should be remains 

elusive. We have quantified storage performance of c. 42,000 

seed accessions, representing 276 species, within the USDA 

National Plant Germplasm System (NPGS) collection, as 

value but great importance to the ecology of high-mountain 

ecosystems of the West. These High Five pines include the 

subalpine and treeline species—whitebark (Pinus albicaulis), 

Rocky Mountain bristlecone (P. aristata), Great Basin bristlecone (P. longaeva), and foxtail (P. balfouriana)—the montane 

to subalpine pine, southwestern white (P. strobiformis), and the 

lower treeline to upper treeline pine, limber (P. flexilis). Here, 

we discuss the taxonomy, distribution, ecology, and Native 

American use of these pines, as well as current threats and 

conservation status. Traditional classification places the bristlecones and foxtail pine together in Subsection Balfourianae, 

limber and southwestern white pine in Subsection Strobi, and 

whitebark pine in Subsection Cembrae. Whitebark pine has 

the largest range and most northerly occurrence. The distribution of limber pine is also large, with a wide elevational range. 

Southwestern white pine occurs from the southwestern U.S. 

through northern Mexico; foxtail pine is found in two widelyseparated regions in California; and, Rocky Mountain bristlecone pine occurs in northern Arizona and the southern Rocky 

Mountains. Great Basin bristlecone pine is restricted to the 

high desert ranges of eastern California, Utah, and Nevada. The 

High Five pines vary successionally and geographically from 

minor to major forest and treeline components. As a group, 

they are also moderately to strongly shade intolerant, and dependent on disturbance, particularly fire, on productive sites 

for forest renewal. The high elevation pines tolerate cold, arid 

sites with poor soils. On exposed sites with infrequent disturbance, these trees can live for 1,000 to 4,500 years, depending 

on the species. Thus, these pines together comprise geographically extensive and ecologically diverse forest habitat types. 

Whitebark, limber and southwestern white pine produce large, 

wingless seeds that are eaten by a diversity of wildlife. Clark’s 

nutcrackers (Nucifraga columbiana) are important seed dispersers for whitebark and limber pine, for southwestern white pine 

in its northern range, and to a lesser extent for the bristlecone 

pines. Furthermore, the High Five pines provide important 

ecosystem services directly benefiting humans, including the 

use of the seeds and other parts of pines as food and medicines 

by Native Americans, the regulation of downstream flow and 

the prevention of soil erosion by treeline forests, and the aesthetic and spiritual values often associated with high elevation 

forests. The future survival of the High Five pines is threatened 

by the exotic blister rust pathogen Cronartium ribicola, current 

mountain pine beetle (Dendroctonus ponderosae) outbreaks, 

successional replacement from fire suppression, and climate 

change. Whitebark pine has been assigned special status in 

Washington and British Columbia, and endangered status 

along with limber pine in Alberta. A petition to list whitebark 

pine as an endangered or threatened species is currently being 

evaluated by the U.S. Fish and Wildlife Service. In Canada, 

whitebark pine has been assessed federally as Endangered and 

is expected to be legally listed soon under the Species at Risk 

Act.

Link: http://www.treesearch.fs.fed.us/pubs/38188
102 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

well as a smaller experiment of 207 cultivars from 42 species. 

Accessions from the NPGS collection were harvested between 

1934 and 1975, and had relatively high initial germination 

percentages that decreased at a variable rate during storage at 

both 5 and -18 °C. Germination time courses, which represent 

the average performance of the species, were fitted to Avrami 

kinetics, to calculate the time at which germination characteristically declined to 50% (P50). These P50 values correlated 

with other longevity surveys reported in the literature for seeds 

stored under controlled conditions, but there was no correlation 

among these studies and seed persistence observed in the classic buried seed experiment by Duvel. Some plant families had 

characteristically short-lived (e.g., Apiaceae and Brassicaceae) 

or long-lived (e.g., Malvaceae and Chenopodiaceae) seeds. 

Also, seeds from species that originated from particular localities had characteristically short (e.g., Europe) or long (e.g., 

South Asia and Australia) shelf lives. However, there appeared 

to be no correlation between longevity and dry matter reserves, 

soluble carbohydrates and parameters relating to soil persistence or resource allocation. Although data from this survey 

support the hypothesis that some species tend to survive longer than others in a genebank environment, there is little information on the attributes of the seed that affect its storage 

performance.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 103

CHAPTER 2 • CONSERVATION AND RESTORATION • STRATEGIES

286. Aubry CA, Rochefort R. 2007. Whitebark pine in 

the Pacific Northwest: what’s next? In: Goheen EM, Sniezko R, 

editors. Whitebark Pine: A Pacific Coast Perspective. Portland 

(OR): USDA Forest Service, Pacific Northwest Region. R6-

NR-FHP-2007-01. p 64-70.

Type: Government Document

Geographic Area: Pacific Northwest USA

Compilers’ Keywords: conservation, decision framework, 

forest management, Pinus albicaulis, restoration

Abstract: Conservation and restoration of whitebark pine are 

two important challenges facing us in the immediate future. 

We will present our approach for a whitebark pine conservation strategy which will focus on Federal lands in Oregon, 

Washington and California. This management plan will identify the actions needed to reduce the threats to the continued 

existence of whitebark pine throughout the planning area. 

Following the completion of the strategy, we will develop a 

land manager’s guide for restoration and managing whitebark 

pine. This manual will describe site assessments, provide a 

decision-making guide for setting stand level prioritization, 

and explain restoration techniques. In our presentation we will 

review these restoration techniques and discuss the challenges 

of their application to the highly variable whitebark pine stands 

in the Pacific Northwest.

Link: http://prdp2fs.ess.usda.gov/Internet/FSE_DOCUMENTS/fsbdev2_026093.pdf

287. Booth DT, Jones TA. 2001. Plants for ecological 

restoration: a foundation and philosophy for the future. Native 

Plants Journal 2:12-20.

Type: Journal 

Geographic Area: Global

Keywords: gene pool, evolution, reclamation, conservation, 

restoration gene pool concept

Abstract: Today’s conservation concerns range from rare 

plant preservation to landscapes threatened by alien annuals. 

Effective action follows effective decision-making based on 

appropriately framed questions. We present the restoration 

gene pool (RGP) concept as a framework for choosing plant 

materials based on the priorities of the species, plant communities, systems, and landscapes threatened. We couch our 

discussion in an acknowledgment of 65 y of national plant materials progress that has evolved with society’s priorities and 

has maintained a high degree of cooperation among participating entities. The plant materials program and its cooperators 

General (215–232)—reviews, surveys, 

summaries, theory, foundations, paradigms

Research (233–285)—propagation, 

outplanting, invasion

Strategies (286–326)—frameworks, approach, 

decision support

Resources (327–338)—native plant materials, 

guidelines, handbooks

Conservation and 

Restoration

#. Author(s), Date. Title. Source.

Type: 

Geographic Area: 

Keywords: 

Description:

Link:

location of study

keywords written by the 

author(s) or provided by the 

compilers

ID number and

information

abstract, summary, or 

portions of the introduction 

or conclusion written by the 

author(s) or provided by the 

compilers

book, book section, conference 

paper, dissertation, government 

document, journal article, 

newsletter, presentation, report, 

thesis or web page

internet address of reference

Each reference contains the following:
104 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

CHAPTER 2 • CONSERVATION AND RESTORATION • STRATEGIES

ture_of_Native_Plants_Used_to_Revegetate_Degraded_Ecosystems/file/32bfe5140b3b283e0d.pdf

289. Davies KW, Boyd CS, Beck JL, Bates JD, 

Svegcar TJ, Gregg MA. 2011. Saving the sagebrush sea: 

an ecosystem conservation plan for big sagebrush plant communities. Biological Conservation 144:2573-2584.

Type: Journal 

Geographic Area: Western USA

Keywords: annual grasses, conservation easements, development, ecosystem management, mitigation, restoration, woodland encroachment

Abstract: Vegetation change and anthropogenic development are altering ecosystems and decreasing biodiversity. 

Successful management of ecosystems threatened by multiple 

stressors requires development of ecosystem conservation 

plans rather than single species plans. We selected the big 

sagebrush (Artemisia tridentata Nutt.) ecosystem to demonstrate this approach. The area occupied by the sagebrush ecosystem is declining and becoming increasingly fragmented 

at an alarming rate because of conifer encroachment, exotic 

annual grass invasion, and anthropogenic development. This 

is causing rangewide declines and localized extirpations of 

sagebrush associated fauna and flora. To develop an ecosystem 

conservation plan, a synthesis of existing knowledge is needed 

to prioritize and direct management and research. Based on 

the synthesis, we concluded that efforts to restore higher elevation conifer-encroached, sagebrush communities were 

frequently successful, while restoration of exotic annual grassinvaded, lower elevation, sagebrush communities often failed. 

Overcoming exotic annual grass invasion is challenging and 

needs additional research to improve the probability of restoration and identify areas where success would be more probable. 

Management of fire regimes will be paramount to conserving 

sagebrush communities, as infrequent fires facilitate conifer 

encroachment and too frequent fires promote exotic annual 

grasses. Anthropogenic development needs to be mitigated 

and reduced to protect sagebrush communities and this probably includes more conservation easements and other incentives to landowners to not develop their properties. Threats to 

the sustainability of sagebrush ecosystem are daunting, but a 

coordinated ecosystem conservation plan that focuses on applying successful practices and research to overcome limitations to conservation is most likely to yield success.

Link: http://oregonstate.edu/dept/eoarc/sites/default/files/

publication/702.pdf

290. Engelmann F. 2012. Germplasm collection, storage, and conservation. Chapter 17. In: Altman A, MichaelHasegawa P, editors. Plant Biotechnology and Agriculture: 

Prospects for the 21st century. London, United Kingdom: 

Academic Press. p 255-267. 

have contributed the bulk of the material and technology now 

used in ecosystem restoration and are foundation for meeting 

conservation challenges of the future. Using a discussion of 

the genetics of native plant materials and 2 conservation challenges, we illustrate how the RGP concept can be used to select 

plant materials based on their ability to meet priority concerns.

Link: http://afrsweb.usda.gov/SP2UserFiles/Place/54281000/

pdfs/tj40.pdf

288. Burton CM, Burton PJ, Hebda R, Turner NJ. 

2006. Determining the optimal sowing density for a mixture of native plants used to revegetate degraded ecosystems.

Restoration Ecology 14:379-390.

Type: Journal 

Geographic Area: Western Canada

Keywords: cover production, degraded soils, ecological restoration, fertilization, seeding densities, soil rehabilitation

Abstract: No standardized, objective methodology exists 

for optimizing seeding rates when establishing herbaceous 

plant cover for pastures, hay fields, ecological restoration, 

or other revegetation activities. Seeding densities, fertilizer 

use, season of seeding, and the interaction of these treatments were tested using native plants on degraded sites 

in northern British Columbia, Canada. A mixture of 20% 

Achillea millefolium, 20% Carex aenea, 20% Elymus glaucus, 

20% Festuca occidentalis, 16% Geum macrophyllum, and 4% 

Lupinus polyphyllus seed was applied at 0, 375, 750, 1,500, 

3,000, and 6,000 pure live seed (PLS) per m2 in 2.5 x 2.5-m 

rototilled test plots, established in the fall and spring, with 

and without fertilizer. There was no significant difference in 

plant cover of sown species between fall seeding and spring 

seeding, and few treatment interactions were identified in 

the first 2 years after sowing. There was no significant difference in cover between seed densities of 3,000 and 6,000 

PLS/m2 in the first year, nor among 1,500, 3,000, and 6,000 

PLS/m2 treatments in the second year. Seed densities as low 

as 375 PLS/m2 produced year 2 plant cover equivalent to 

that observed at 3,000 PLS/m2 in year 1. Plots sown to seed 

densities less than or equal to 750 PLS/m2 generally exhibited an increase (infilling) in plant density from year 1 to year 

2, whereas plots sown to seed densities greater than or equal 

to 1,500 PLS/m2 generally exhibited a decrease (densitydependent mortality) in plant density. These results imply a 

most efficient sowing density between 750 and 1,500 PLS/

m2 (corresponding to 190–301 established plants m-2 after 

two growing seasons). It is suggested that net changes in 

plant populations observed over a range of sowing densities are a robust and objective means of determining optimal sowing densities for the establishment of herbaceous 

perennials.

Link: http://www.researchgate.net/publication/229671456_

Determining_the_Optimal_Sowing_Density_for_a_Mix-
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 105

Geographic Area: Global

Compilers’ Keywords: ecological function, approaches, climate change, paleoecology, framework

Introduction: This book is meant to provide a scientific 

framework for restoration ecology that can be used to inform 

ecological restoration as well as stimulate advances in our understanding of nature. As you read, bear in mind that the implementation of ecological restoration is not only escalating at an 

astounding rate, but also that it remains the most ecologically 

viable and aesthetically appealing remedy for mending Earth’s 

ever increasing number and scale of degraded ecosystems.

Link: http://xa.yimg.com/kq/groups/22969879/472658511/

name/Foundations+of+Restoration+Ecology.pdf#page=328

293. Ford-Lloyd BV, Jackson MT. 1991. Biotechnology 

and methods of conservation of plant genetic resources. Journal 

of Biotechnology 17:247-256.

Type: Journal 

Geographic Area: Global

Keywords: genetic diversity, germplasm, conservation, genebank, cryopreservation

Introduction: The loss of biological diversity is of major environmental concern at the present time. Whether this relates 

to the destruction of the tropical rain forests, for instance, or 

the threat which faces natural ecosystems as climates change 

due to global warming, it is important that in order to survive 

and continue to evolve, plant and animal species must contain an ample reservoir of genetic diversity. One aspect of this 

process is the continuing loss of genetic diversity in the crops 

upon which world agriculture is based. Scientists throughout 

the world are rightly engaged in developing better and higher 

yielding cultivars of crop plants to be used on increasingly 

larger scales. But this involves the replacement of the generally variable, lower yielding, locally adapted strains grown in 

traditional farming systems, by the products of modern agriculture. Nowadays every major crop has a relatively narrow 

genetic base, and so under such situations, diversity in farming 

systems is replaced by genetic uniformity. However, in order 

to be able to respond to the various stresses that threaten modern agriculture, plant breeders are dependent upon the availability of a pool of diverse genetic material. The significance 

of genetic uniformity can be highlighted by several crop examples, one of historical importance and two others of more 

recent times. The history of the potato in Europe illustrates the 

necessity for utilizing genetic resources to broaden the genetic 

base of crop plants. The narrow genetic base of the potato crop 

was recognised during the last century following the devastating epidemics of late blight disease, caused by Phytophthora 

infestans, in the 1840s in Ireland. Massive destruction of the 

corn crop came about in the United States in 1970 with the 

southern corn leaf blight epidemic (caused by the fungus 

Helmmthosportum maydls). Modern plant breeding had led to 

Type: Book Section

Geographic Area: Global

Compilers’ Keywords: biodiversity, ex situ, in situ, in vitro, cryopreservation, orthodox seeds, recalcitrant seeds, seed 

banking, climate change, genetic loss, gene conservation

Introduction: Two basic conservation strategies, in situ and 

ex situ, each composed of various techniques, are employed 

for conservation of plant biodiversity. Until recently, most conservation efforts, apart from work on forest genetic resources, 

have concentrated on ex situ conservation, particularly seed 

genebanks. In the 1950s and 1960s, major advances in plant 

breeding brought about the Green Revolution, which resulted 

in the wide-scale adoption of high-yielding varieties and genetically uniform cultivars of staple crops, particularly wheat 

and rice. Consequently, global concern about the loss of genetic diversity in these crops increased, as farmers abandoned 

their locally adapted landraces and traditional varieties and 

replaced them with improved, yet genetically uniform, modern 

ones. In response to this concern, research groups started to assemble germplasm collections of the major crop species within 

their respective mandates. It is in this context that the IBPBR 

(Biodiversity International) was established in 1974 to coordinate the global effort to systematically collect and conserve the 

world’s threatened plant genetic diversity.

291. Falk DA, Knapp EE, Guerrant EO Jr. 2001. An 

introduction to restoration genetics. Tucson (AZ): Society for 

Ecological Restoration. 33 p.

Type: Report

Geographic Area: Global

Compilers’ Keywords: climate change, diversity, review, 

guide, ecological restoration, policy, adaptation, uncertainty

Introduction: All living organisms carry a genetic blueprint. 

This is so regardless of whether they are plants, animals, or 

fungi, whether they are short- or long-lived, and whether they 

reproduce sexually or clonally. Therefore, to the extent that 

restoration deals with living organisms, genetics are part of the 

picture. Although the basic principles underlying restoration 

genetics may be familiar, to date surprisingly little attention 

has been devoted to genetic considerations in restoration practice. The purpose of this Restoration Science and Policy Paper 

is to outline some considerations that restoration designers and 

managers should be aware of, and to identify more detailed 

resources that may be useful in practice.

Link: http://www.ser.org/docs/default-document-library/

ser_restoration_genetics.pdf

292. Falk DA, Palmer MA, Zedler JB. 2006.

Foundations of restoration ecology. Washington (DC): Island 

Press. 379 p. 

Type: Book

CHAPTER 2 • CONSERVATION AND RESTORATION • STRATEGIES
106 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

genetic erosion and gene flow. A comparison of past and present genecological distributions (patterns of genetic variation 

of key adaptive traits in the ecological space) of selected species is a realistic way of assessing the trend of intra-specific 

variation, and thus provides a state indicator of tree genetic 

diversity also able to reflect possible pressures threatening genetic diversity. Revealing benefits of genetic diversity related 

to ecosystem services is complex, but current trends in plantation performance offer the possibility of an indicator of benefit. Response indicators are generally much easier to define, 

because recognition and even quantification of, e.g., research, 

education, breeding, conservation, and regulation actions and 

programs are relatively straightforward. Only state indicators 

can reveal genetic patterns and processes, which are fundamental for maintaining genetic diversity. Indirect indicators 

of pressure, benefit, or response should therefore not be used 

independently of state indicators. A coherent set of indicators 

covering diversity–productivity–knowledge–management 

based on the genecological approach is proposed for application on appropriate groups of tree species in the wild and in 

cultivation worldwide. These indicators realistically reflect the 

state, trends and potentials of the world’s tree genetic resources 

to support sustainable growth. The state of the genetic diversity 

will be based on trends in species population distribution and 

diversity patterns for selected species. The productivity of the 

genetic resource of trees in current use will reflect the possible 

potential of mobilizing the resource further. Trends in knowledge will underpin the potential capacity for development of 

the resource and current management of the genetic resource 

itself will reveal how well we are actually doing and where 

improvements are required.

Link: http://www.sciencedirect.com/science/article/pii/

S0378112714002825

295. Hargrove WM, Hoffman FM. 2005. Potential of 

multivariate quantitative methods for delineation and visualization of ecoregions. Environmental Management 34:S39-S60.

Type: Journal 

Geographic Area: USA

Compilers’ Keywords: clustering, climate change, ecotone, 

environmental envelope, niche, preserve design, time series

Abstract: Multivariate clustering based on fine spatial resolution maps of elevation, temperature, precipitation, soil characteristics, and solar inputs has been used at several specified 

levels of division to produce a spectrum of quantitative ecoregion maps for the conterminous United States. The coarse 

ecoregion divisions accurately capture intuitively-understood 

regional environmental differences, whereas the finer divisions 

highlight local condition gradients, ecotones, and clines. Such 

statistically generated ecoregions can be produced based on 

user-selected continuous variables, allowing customized regions to be delineated for any specific problem. By creating 

the use of a cytoplasmic gene which conferred susceptibility 

to a particular race of this fungal pathogen. A more recent, but 

less publicised case of genetic vulnerability was caused not 

by plant disease, but by cold weather. By 1972 in the Soviet 

Union the wheat variety ‘Bezostaja’ was grown on almost 15 

million hectares. It had been moved beyond its original area of 

cultivation into the Ukraine during a period of relatively mild 

winters. Then in 1972 a very severe winter occurred, causing 

losses of millions of tons of winter wheat (Fischbeck, 1981). In 

relation to this example, we can ask whether global warming 

will lead to similar crop losses in the future (Jackson and FordLloyd, 1990). Plant genetic resources (germplasm) is a term 

used to describe the total genetic diversity of cultivated species 

and their wild relatives, much of which may be of value to 

breeders. Although commercial and obsolete varieties, breeders’ lines and induced or natural mutations may be included 

under this term, it is much more usual to describe plant genetic 

resources as landraces or primitive forms, weed races which 

are closely related to the cultivated species, and related wild 

species. Landraces are populations of crops, often collected 

from remote areas, where the new, highly bred cultivars have 

not been introduced. They are highly diverse genetically, and 

have often been grown as mixtures of species, as well as diverse populations of one species. Such materials are closely 

allied genetically to modern varieties and are extremely important genetic resources, and as such have received top priority 

for conservation.

294. Graudal L, Aravanopoulos F, Bennadji 

Z, Changtragoon S, Fady B, Kjær ED, Loo J, 

Ramamonjisoa L, Vendramin GG. 2014. Global 

to local genetic diversity indicators of evolutionary potential 

in tree species within and outside forests. Forest Ecology and 

Management 333:35-51.

Type: Journal 

Geographic Area: Global

Keywords: types of indicators, genetic and demographic 

verifiers, the genecological approach, diversity-productivityknowledge management

Abstract: There is a general trend of biodiversity loss at 

global, regional, national and local levels. To monitor this 

trend, international policy processes have created a wealth 

of indicators over the last two decades. However, genetic diversity indicators are regrettably absent from comprehensive 

bio-monitoring schemes. Here, we provide a review and an 

assessment of the different attempts made to provide such indicators for tree genetic diversity from the global level down to 

the level of the management unit. So far, no generally accepted 

indicators have been provided as international standards, nor 

tested for their possible use in practice. We suggest that indicators for monitoring genetic diversity and dynamics should 

be based on ecological and demographic surrogates of adaptive diversity as well as genetic markers capable of identifying 

CHAPTER 2 • CONSERVATION AND RESTORATION • STRATEGIES
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 107

CHAPTER 2 • CONSERVATION AND RESTORATION • STRATEGIES

Type: Journal 

Geographic Area: Global

Keywords: genetic adaptation, genetic identity, metapopulation, multiple-origin polycross

Abstract: Restoration practitioners have long been faced 

with a dichotomous choice of native versus introduced plant 

material confounded by a general lack of consensus concerning what constitutes being native. The “restoration gene pool” 

concept assigns plant materials to one of four restoration gene 

pools (primary to quaternary) in order of declining genetic correspondence to the target population. Adaptation is decoupled 

from genetic identity because they often do not correspond, 

particularly if ecosystem function of the disturbed site has been 

altered. Because use of plant material with highest genetic 

identity, that is, the primary restoration gene pool, may not be 

ultimately successful, material of higher order pools may be 

substituted. This decision can be made individually for each 

plant species in the restored plant community in the scientific 

context that ecosystem management demands. The restoration 

gene pool concept provides a place for cultivars of native species and noninvasive introduced plant material when use of 

native-site material is not feasible. The use of metapopulation 

polycrosses or composites and multiple-origin polycrosses or 

composites is encouraged as appropriate. The restoration gene 

pool concept can be implemented as a hierarchical decisionsupport tool within the larger context of planning seedings.

Link: http://www.fs.fed.us/rm/boise/research/shrub/

Links/2003papers/jones2003.pdf

298. Jones TA, Johnson DA. 1998. Integrating genetic 

concepts into planning rangeland seedings. Journal of Range 

Management 51:594-606.

Type: Journal 

Geographic Area: USA

Keywords: genetic integrity, genetic pollution, introgression, 

natural selection, plant materials, polyploidy, rangeland weeds, 

succession

Abstract: Choice of plant materials is a fundamental component of any rangeland rehabilitation, reclamation, or restoration 

project. We describe here an integrated approach for such decision-making. This approach considers site potential, desired 

landscape, seeding objectives, conflicting land use philosophies, appropriate plant materials, weed invasion, community 

seral status, and economic limitations. Technical limitations 

are considered in generating a plan that has the greatest potential for success. Determining whether native-site plant material is best depends on objectives, heterogeneity of the site’s 

environment, uniqueness of the site, plant population size, and 

biotic or abiotic site disturbance. Fixation of alien genes into 

a population is referred to both as introgression, which may 

ensure maintenance of genetic variation critical for adaptation 

an objective ecoregion classification, the ecoregion concept is 

removed from the limitations of human subjectivity, making 

possible a new array of ecologically useful derivative products. 

A red–green–blue visualization based on principal components 

analysis of ecoregion centroids indicates with color the relative 

combination of environmental conditions found within each 

ecoregion. Multiple geographic areas can be classified into 

a single common set of quantitative ecoregions to provide a 

basis for comparison, or maps of a single area through time 

can be classified to portray climatic or environmental changes 

geographically in terms of current conditions. Quantified representativeness can characterize borders between ecoregions 

as gradual, sharp, or of changing character along their length. 

Similarity of any ecoregion to all other ecoregions can be 

quantified and displayed as a ‘‘representativeness’’ map. The 

representativeness of an existing spatial array of sample locations or study sites can be mapped relative to a set of quantitative ecoregions, suggesting locations for additional samples or 

sites. In addition, the shape of Hutchinsonian niches in environment space can be defined if a multivariate range map of 

species occurrence is available.

Link: http://www.researchgate.net/publication/7856972_Potential_of_multivariate_quantitative_methods_for_delineation_and_visualization_of_ecoregions/file/d912f508063c35b761.pdf

296. Havens K, Vitt P, Maunder M, Guerrant EO 

Jr, Dixon KW. 2006. Ex situ plant conservation and beyond.

BioScience 56:525-531.

Type: Journal 

Geographic Area: USA

Keywords: botanic garden, performance indicators, conservation strategies

Abstract: In recent years, the mission of many botanic gardens and arboreta has expanded from a traditional focus on 

developing a horticultural collection to one that includes taking a proactive role in plant conservation. To use their limited 

resources more effectively, many gardens are seeking ways to 

quantify their contributions to conservation efforts, both as a 

self-assessment tool to improve their effectiveness and as a 

way to give an explicit accounting of activities to donors and 

funding agencies. We suggest many ways gardens can measure 

the success of their conservation programs, and present results 

from a survey conducted to assess current conservation activities at botanic gardens.

Link: http://www.researchgate.net/publication/232691529_Ex_Situ_Plant_Conservation_and_Beyond/

file/5046351a4cde1f01e9.pdf

297. Jones TA. 2003.The restoration gene pool concept: beyond the native versus non-native debate. Restoration Ecology 

11:281-290.
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CHAPTER 2 • CONSERVATION AND RESTORATION • STRATEGIES

RE, Tomback DF, Murray MP, Smith CM, editors. High Five 

Symposium. Fort Collins (CO): USDA Forest Service, Rocky 

Mountain Research Station. Proceedings RMRS-P-63. p 

276-294.

Type: Government Document

Geographic Area: Western North America

Compilers’ Keywords: conference paper, white pine forests, 

mountain pine beetle, Dendroctonus spp., conservation, keystone species, restoration, strategies

Abstract: Many ecologically important, five-needle white 

pine forests that historically dominated the high elevation landscapes of western North America are now being heavily impacted by mountain pine beetle (Dendroctonus spp.) outbreaks, 

the exotic disease white pine blister rust (WPBR), and altered 

high elevation fire regimes. Management intervention using 

specially designed strategic treatments will be needed to conserve these keystone species. The goal of this intervention is to 

promote self-sustaining five-needle white pine ecosystems that 

have both resilience to disturbances and genetic resistance to 

white pine blister rust. Many tools and methods are available 

for land managers. In this paper we outline important steps for 

implementation of restoration treatments in declining high elevation white pine stands and discuss a number of proactive 

treatments in threatened ecosystems to mitigate adverse impacts of rust, beetles, and lack of fire. These steps for restoration include (1) create a strategy for restoration across multiple 

scales, (2) develop materials and techniques for conducting 

restoration treatments, such as seed collections and rust resistance assessments of the genetic material, (3) prioritize stands 

or landscapes by integrating the strategy with other management issues, administrative barriers, climate change mitigation, and other local concerns, (4) implement silvicultural cuttings and prescribed fire according to landscape and stand level 

strategies, (5) conduct activities and assessments to enhance 

and ensure restoration treatments are effective including planting rust-resistant pine seedlings and protecting valuable seedsources, and finally (6) monitor treated landscape and stands 

for effects and adjust and modify future treatment designs accordingly. Examples from whitebark pine ecosystems in the 

northern Rocky Mountains will be presented to demonstrate 

this process. For those high elevation white pine ecosystems 

that are threatened by white pine blister rust or mountain pine 

beetle, there are actions that can be taken proactively to gain 

necessary information to evaluate risk and prepared landscapes 

for invasion to mitigate future impacts. The proactive strategy 

includes: (1) educate and engage the public and managers to 

shift from crisis management to management for resiliency, (2) 

conserve genetic diversity from native populations before they 

are impacted by WPBR or other stresses, (3) conduct research 

on patterns, processes and responses of native ecosystems to 

provide process level understanding of ecosystem behavior 

and (4) develop and conduct appropriate management activities to increase the resiliency of high elevation five-needle pine 

to a changing environment, and as genetic pollution, with the 

potential for swamping native cross-pollinating annual or 

short-lived perennial gene pools. Precautionary procedures 

during seed increase minimize genetic shift, which may be 

reversible, but genetic drift could result in permanent loss of 

desirable genes. A variety of germplasm classes, ranging from 

site-specific to widely adapted and varying in degrees of heterozygosity and heterogeneity should be considered. Material 

originating from multiple sites may increase the opportunity 

for natural selection. An understanding of the magnitude and 

nature of a species’ genetic variation, its relationship to ecological adaptation, and its interaction with other ecosystem 

components contribute to informed decision-making. Though 

often unavailable, experience is the best guide for predicting 

performance of materials on non-native sites. 

Link: http://afrsweb.usda.gov/SP2UserFiles/Place/54281000/

pdfs/tj33.pdf

299. Jones TA, Monaco TA. 2007. A restoration practitioner’s guide to the restoration gene pool concept. Ecological 

Restoration 25:12-19.

Type: Journal 

Geographic Area: USA

Keywords: plant materials, genetic identity, metapopulation, 

ecoregion, restoration gene pool concept, ecological restoration

Abstract: Choosing plant materials for each desired species 

is often one of the most difficult steps in developing a restoration plan. The Restoration Gene Pool concept was developed 

to clarify the options available to the ecological restoration 

practitioner in terms of plant materials. We present a decisionmaking flowchart incorporating the issues delineated in the 

Restoration Gene Pool concept. We intend to provide practitioners with a framework to make objective and defensible plant 

materials choices in keeping with the objectives and philosophy of the restoration project. The flowchart consists of a series 

of boxes with single or multiple statements to be judged as true 

or false. The statements deal with issues such as genetic identity, plant metapopulation, functional guild, ecoregion, invasive 

weed presence, site soil and climate, endangered species presence, commercial seed availability, seed contracting potential, 

and availability of material bred for improved stress tolerance. 

Implementing the flowchart is a straightforward way to apply 

the Restoration Gene Pool concept to a particular project, but 

we encourage each practitioner to personalize the flowchart to 

make it as practical as possible for the situations that she or he 

most often encounters.

Link: http://www.fs.fed.us/rm/pubs_other/rmrs_2007_jones_

t001.pdf

300. Keane RE, Schoettle AW. 2011. Strategies, tools, 

and challenges for sustaining and restoring high elevation fiveneedle white pine forests in western North America. In: Keane 
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 109

CHAPTER 2 • CONSERVATION AND RESTORATION • STRATEGIES

seed transfer in ponderosa pine and Douglas-fir. Genetic gains 

in adaptation, and insect and disease resistance, continue to be 

made in western white pine, western larch, ponderosa pine, 

Douglas-fir, and lodgepole pine. While progress has been made 

in the white pine blister rust program, restoring western white 

pine to Inland Northwest forests requires a continued commitment to selective breeding. Other insect and disease problems 

should also receive strong consideration in selective breeding 

programs, to prevent erosion of existing genetic resistance, and 

when warranted, to sustain and enhance this resistance.

Link: http://www.treesearch.fs.fed.us/pubs/23502

303. McArthur ED, Young SA. 1999. Development of 

native seed supplies to support restoration of piñon-juniper sites. 

In: Monsen SB, Stevens R, editors. Ecology and Management 

of Piñon-Juniper Communities within the Interior West. Ogden 

(UT): USDA Forest Service, Rocky Mountain Research 

Station. Proceedings RMRS-P-9. p 327-330.

Type: Government Document

Geographic Area: Western USA

Compilers’ Keywords: woodlands, piñon-juniper, land management, survey, native plant materials

Abstract: Use of native plants for restoration and rehabilitation of disturbed or manipulated piñon-juniper communities is 

increasing in response to desires of land managers and society 

in general. Seeds of native plants are becoming more available, 

but estimates and surveys show there is still more demand than 

available supply. Field grown seeds and warehousing do anticipate demand but are only partial solutions to the native plant 

seed shortage. Exotic, developed plant materials, especially 

Triticeae grasses and legumes, remain important resources for 

rehabilitation plantings. Private industry seed collectors, growers, and developers will be responsive to plant materials needs 

of land managers. Seed genetic identity and quality can be better assured through the seed certification process whether the 

seed is wildland collected or field grown.

Link: http://www.fs.fed.us/rm/pubs/rmrs_p009/rmrs_

p009_327_330.pdf

304. McMahon G, Gregonis SM, Waltman SW, 

Omernik JM, Thorson TD, Freeouf JA, Rorick AH, 

Keys JE. 2001. Developing a spatial framework of common 

ecological regions for the conterminous US. Environmental 

Management 28:293-316.

Type: Journal 

Geographic Area: USA

Keywords: ecological region, spatial framework, ecologically 

oriented management, land resources, ecosystem management

Abstract: In 1996, nine federal agencies with mandates to 

inventory and manage the nation’s land, water, and biological resources signed a memorandum of understanding entitled 

ecosystems to prepare them for change. Whether it is restoring 

impacted landscapes or interventions to mitigate the development of impacts on threatened landscapes, there are two important factors that will govern the success of these species 

even with comprehensive and effective rangewide strategies: 

(1) the magnitude of resources available over time to conduct 

restoration efforts, and (2) the commitment of natural resource 

agencies to conduct restoration activities over the long term, 

most likely for many decades to centuries.

Link: http://www.treesearch.fs.fed.us/pubs/38240

301. MacKinnon A, Meidinger DV, Klinka K. 1992.

Use of the biogeoclimatic ecosystem classification in British 

Columbia. The Forestry Chronicle 68:100-120.

Type: Journal 

Geographic Area: Western Canada

Compilers’ Keywords: silviculture, land management, land 

classification

Abstract: Biogeoclimatic ecosystem classification (BEC) is a 

hierarchical system that organizes ecosystems at three levels of 

integration - local, regional, and chronological. The system is 

used by silviculturists and range, recreation and wildlife managers in British Columbia. Based on the study of both vegetation and sites, the system reveals ecological potentialities and 

limitations of particular sites, and combined with the accumulation and widespread dissemination of ecological knowledge 

provides an ideal framework for integrated resource management. The ways in which the BEC system is adapted for, and 

used by, resource managers are described and demonstrated. 

Ongoing activities, including classification of seral ecosystems, quantification of soil moisture and nutrient regimes, 

identification of relationships between forest productivity and 

site quality, and provincial correlation of the six regional classifications, are outlined. 

302. Mahalovich MF. 1995. The role of genetics in improving forest health. In: Eskew LG, editors. 1995 National 

Silviculture Workshop: Forest Health through Silviculture. 

Fort Collins (CO): USDA Forest Service, Rocky Mountain 

Forest and Range Experiment Station. General Technical 

Report RM-GTR-267. p 200-207.

Type: Government Document

Geographic Area: Northwestern USA

Compilers’ Keywords: forest management, seed transfer, 

guidelines

Abstract: An often ignored tool to improve forest health is 

the application of genetics. Tree improvement programs in the 

Inland West utilize genetic principles to develop-seed transfer 

guidelines to avoid the problems associated with off-site plantings and to improve characteristics in conifers related to forest 

health. PC-based expert systems have been developed to aid in 
110 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

CHAPTER 2 • CONSERVATION AND RESTORATION • STRATEGIES

Link: http://www.internationalgrasslands.org/files/igc/publications/1997/iii-299.pdf

306. Millar CI. 2014. Historic variability: informing restoration strategies, not prescribing targets. Journal of Sustainable 

Forestry 33:S28-S42.

Type: Journal 

Geographic Area: North America

Keywords: climate change, ecological restoration, ecosystem 

management, forest management, historical ecology, historic 

range of variability

Abstract: The concept of historic range of variability (HRV) 

is briefly evaluated within the context of its application in 

ecosystem management over the past two decades. Despite 

caveats to the contrary, an implicit assumption continues to 

emerge of climatic stationarity, and, by corollary, that presettlement centuries provide an appropriate reference period. 

This is examined from the perspective of historic climate 

change and ecosystem response. As a means of developing 

reference prescriptions and management targets, HRV is 

generally inappropriate, although if historic periods are used 

for reconstruction that have coarse resemblance to present 

or projected future climates, such as the Medieval Climate 

Anomaly or middle Holocene rather than the pre-settlement 

centuries, these might be defensible. In cases of reclamation 

of severely degraded ecosystems, HRV prescriptions developed from analogous climate periods could provide coarse 

guides. In most situations, however, historic reconstructions are best used to improve understanding of ecological 

response to a wide range of forcing factors, and thereby to 

inform (rather than prescribe) management strategies. Such 

historically informed approaches are likely more effective 

than an HRV approach under future changing climate regimes for managing and restoring ecosystem function and 

for assisting transitions to new ecosystem states.

Link: http://www.treesearch.fs.fed.us/pubs/47361

307. Neale DB, Kremer A. 2011. Forest tree genomics: 

growing resources and applications. Nature 12:111-122.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: genomics, climate change

Abstract: Over the past two decades, research in forest tree 

genomics has lagged behind that of model and agricultural 

systems. However, genomic research in forest trees is poised 

to enter into an important and productive phase owing to the 

advent of next-generation sequencing technologies, the enormous genetic diversity in forest trees and the need to mitigate 

the effects of climate change. Research on long-lived woody 

perennials is extending our molecular knowledge of complex 

life histories and adaptations to the environment—enriching a 

“Developing a Spatial Framework of Ecological Units of the 

United States.” This spatial framework is the basis for interagency coordination and collaboration in the development of 

ecosystem management strategies. One of the objectives in 

this memorandum is the development of a map of common 

ecological regions for the conterminous United States. The 

regions defined in the spatial framework will be areas within 

which biotic, abiotic, terrestrial, and aquatic capacities and potentials are similar. The agencies agreed to begin by exploring 

areas of agreement and disagreement in three federal naturalresource spatial frameworks—Major Land Resource Areas of 

the U.S. Department of Agriculture (USDA) Natural Resources 

Conservation Service, National Hierarchy of Ecological Units 

of the USDA Forest Service, and Level III Ecoregions of the 

U.S. Environmental Protection Agency. The explicit intention 

is that the framework will foster an ecological understanding of the landscape, rather than an understanding based on a 

single resource, single discipline, or single agency perspective. 

This paper describes the origin, capabilities, and limitations of 

three major federal agency frameworks and suggests why a 

common ecological framework is desirable. The scientific and 

programmatic benefits of common ecological regions are described, and a proposed process for development of the common framework is presented.

305. Meyer SE. 1999.Genecological considerations in grassland restoration using wild-collected seed sources. In: BuchananSmith JG, Bailey LD, McCaughey P, editors. Proceedings of 

the 18th International Grassland Congress. Calgary, Alberta: 

Association Management Center. p 299-304.

Type: Conference Paper

Geographic Area: Western USA

Keywords: disturbed land reclamation, ecological restoration, 

genetic variation, propagation, seed transfer guidelines

Abstract: Choice of species and seed sources to use in the 

revegetation of drastically disturbed lands depends on intended post-reclamation land use. When the intent is to return the 

land to predisturbance native plant cover and composition, the 

process can be termed ecological restoration. Seed sources for 

species used in restoration are usually wildland populations. 

Genecology deals with the ecological or adaptive significance 

of genetic variation within and among populations of plants. In 

contrast with plant breeding or biotechnological approaches, 

in restoration this knowledge is used to safeguard the genetic 

composition of source populations, restored populations, and 

populations adjacent to restored areas. Particular attention is 

paid to adequate sampling of source populations and to the 

maintenance of within-population genetic diversity through 

seed increase or nursery propagation phases. The goal is to 

initiate processes that will lead to the reestablishment of native 

ecosystems with diversity, resilience, and capacity for continued evolution, nor to try to create static, idealized predisturbance vegetation.
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 111

Abstract: In this paper, we propose the ecological network 

concept as a suitable basis for inserting biodiversity conservation into sustainable landscape development. For landscapes 

to be ecologically sustainable, the landscape structure should 

support those ecological processes required for the landscape 

to deliver biodiversity services for present and future generations. We first show that in multifunctional, human dominated 

landscapes, biodiversity conservation needs a coherent largescale spatial structure of ecosystems. Theory and empirical 

knowledge of ecological networks provides a framework for 

the design of such structures. Secondly, ecological networks 

can bridge the paradox between reserve conservation (fixing 

nature in space and time) and development, which implies 

change. This is because ecological networks can change 

structure without losing their conservation potential. Thirdly, 

ecological networks facilitate stakeholder decision-making on 

feasible biodiversity goals. They help to focus on an effective 

spatial scale. We conclude that extending the ecological network concept with multifunctional indicators is a promising 

step towards sustainable landscape development and stakeholder decision-making.

310. Perry N. 2010. The ecological importance of species 

and the Noah’s Ark problem. Ecological Economics 69:478-485.

Type: Journal 

Geographic Area: Global

Keywords: ecological importance, ecosystem process, functional diversity, biodiversity, endangered species conservation

Abstract: Using the Noah’s Ark problem—the problem of efficiently allocating limited funds to conserve biodiversity - the 

standard economic approach to endangered species conservation constructs a human-centered biodiversity by favoring species directly valuable to humans. I analyze this approach and 

draw on the functional ecology literature to offer an alternative 

emphasizing the role species play in their ecosystems. The aim 

is to create a working ecosystem on the Ark rather than a collection of charismatic and distinct species. To do so, I construct 

a new measure of a species’ ecological importance and an 

ecological objective appropriate for cost-effective resource allocation. The ecological approach fundamentally changes the 

notion of species-value from a direct value based on a species’ 

appearance or taxonomic difference to an indirect value based 

on a species’ ecological role in its ecosystem. In the process, 

‘populations’ of species become the fundamental unit of biodiversity rather than ‘species,’ and biotic processes also possess 

value. When compared to the economic approach, the ecological approach prioritizes different species for the Ark and 

achieves superior economic outcomes in all but the mythical 

Noah’s Ark scenario where interactions are non-existent. The 

analysis challenges the approach of U.S. endangered species 

legislation and I call for a reformulation based on endangered 

ecological interactions. 

field that has traditionally drawn biological inference from a 

few short-lived herbaceous species.

Link: http://www.ncbi.nlm.nih.gov/pubmed/21245829

308. Nuismer SL, Gandon S. 2008. Moving beyond 

common-garden and transplant designs: insight into the causes 

of local adaptation in species interactions. The American 

Naturalist 171:658-668.

Type: Journal 

Geographic Area: Global

Keywords: coevolution, gene flow, geographic mosaic, hostparasite, Red Queen

Abstract: Theoretical and empirical studies of local adaptation in species interactions have increased greatly over the 

past decade, yielding new insights into the conditions that 

favor local adaptation or maladaptation. Generalizing the results of these studies is difficult, however, because of the different experimental designs that have been used to infer local adaptation. Particularly challenging is comparing results 

across empirical studies conducted in a common laboratory 

or garden environment with results of those conducted using transplants in natural environments. Here we develop 

simple and easily interpretable mathematical expressions for 

the quantities measured by these two different types of studies. Our results reveal that common-garden designs measure 

only a single component of local adaptation-the spatial covariance between the genotype frequencies of the interacting 

species-and thus provide only a partial description of local 

adaptation. In contrast, reciprocal-transplant designs incorporate additional terms that measure the contribution of spatial variability in the ecological environment. Consequently, 

the two types of studies should yield identical results only 

when local adaptation is caused by spatial variability in the 

genotype frequencies of the interacting species alone. In order to unify these disparate approaches, we develop a new 

methodology that can be used to estimate the individual 

components of local adaptation. When implemented in an 

appropriate experimental system, this partitioning allows the 

examination of fundamental questions such as the relative 

proportion of local adaptation attributable to interactions between species or to the abiotic environment.

Link: http://www.ncbi.nlm.nih.gov/pubmed/18419564

309. Opdam P, Steingröver E, van Rooij S. 2006.

Ecological networks: a spatial concept for multi-actor planning 

of sustainable landscapes. Landscape and Urban Planning 

75:322-332.

Type: Journal 

Geographic Area: Global

Keywords: ecological networks, landscape planning, stakeholder decision-making, biodiversity, sustainable development

CHAPTER 2 • CONSERVATION AND RESTORATION • STRATEGIES
112 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Link: http://www.ncbi.nlm.nih.gov/pubmed/20618901

313. Richardson BA, Ekramoddoulah AKM, Liu JJ, 

Kim M-S, Klopfenstein NB. 2010. Current and future 

molecular approaches to investigate the white pine blister rust 

pathosystem. Forest Pathology 40:314-331.

Type: Journal 

Geographic Area: Western USA

Compilers’ Keywords: Pinus, genetic variation

Abstract: Molecular genetics is proving to be especially useful for addressing a wide variety of research and management 

questions on the white pine blister rust pathosystem. White 

pine blister rust, caused by Cronartium ribicola, is an ideal 

model for studying biogeography, genetics, and evolution because: (1) it involves an introduced pathogen; (2) it includes 

multiple primary and alternate hosts occurring in large, relatively undisturbed ecosystems; (3) some hosts exhibit endemic 

resistance; and (4) the disease interaction is long enduring. 

Molecular techniques are used to investigate population genetics, phylogenetics, hybrids, and proteomics in white pine 

(Pinus, subgenus Strobus) and blister rust (Cronartium) and the 

genetics of resistance and virulence in the blister rust pathosystem. These techniques include genetic markers, mapping, 

microarrays, sequencing, association genetics, genomics, and 

genecology. Molecular genetics contributes to gene conservation, breeding for resistance, and ecosystem management.

Link: http://www.treesearch.fs.fed.us/pubs/36219

314. Riggs LA. 1990. Conserving genetic resources onsite in forest ecosystems. Forest Ecology and Management 

35:45-68.

Type: Journal 

Geographic Area: California, USA

Compilers’ Keywords: in situ, forest management, decision 

framework, genetic diversity, Douglas-fir, Pseudotsuga menziesii, ecosystem services, genetic conservation

Abstract: Genetic diversity and its structure (its organization 

in space and time) are the critical raw materials from which 

many other aspects of diversity are derived. These genetic 

resources represent information about unique and successful 

relationships among genes and between gene complexes and 

environments. Only a fraction of this information has been 

mined through research. The balance, and the genetic materials themselves, are most effectively conserved for future use 

by on-site (in-situ) preservation, management, or restoration 

of populations, communities and entire landscapes. While 

we have far to go on the road to understanding biodiversity, 

research progressing in an array of disciplines offers ample 

justification to maintain evolved patterns and processes that 

underlie biodiversity, mediate the efficiency of ecosystem ‘services’, and influence the availability of genetic raw material 

311. Pojar J, Klinka K, Meidinger DV. 1987.

Biogeoclimatic ecosystem classification in British Columbia.

Forest Ecology and Management 22:119-154.

Type: Journal 

Geographic Area: Western Canada

Compilers’ Keywords: land classification, land management, forest management

Abstract: Biogeoclimatic ecosystem classification is a system 

of ecological classification widely used in British Columbia. 

The system has been expanded by the B.C. Forest Service 

from the pioneering work of V.J. Krajina and his students. 

The recognized units result from a synthesis of vegetation, 

climate, and soil data. The approach to classification is hierarchical with three interrelated levels of integration. The 

multiple-category, local and regional levels involve vegetation 

and site; a chronological level deals with vegetation dynamics. 

Taxa of any level and category can be integrated according to 

their interpretive value for management. Vegetation units are 

defined and arranged into a floristic hierarchy based on the 

plant association, using traditional Braun-Blanquet methods 

modified to suit regional conditions. Where possible, zonal 

(climatic climax) plant associations are identified, thus defining biogeoclimatic subzones, which are divided into variants 

and aggregated into zones, regions, and formations. Plant associations are transformed into site associations environmentally characterized classes of ecosystems with similar biotic 

potential. Site associations are then subdivided into series and 

types, using biogeoclimatic and edaphic characteristics as differentiae. Examples of classification at local and regional levels are provided, as are examples of management applications. 

Biogeoclimatic ecosystem classification is a unique approach 

that draws on several of the European (including Russian) and 

North American schools of vegetation and land classification, 

and has similarities with the Cajander forest type, Barnes land 

type, Daubenmire habitat type, and Hills site type approaches. 

The classification provides a powerful integrative and predictive tool, with proven practical value for forest managers.

312. Renn SC, Siemens DH. 2010. Ecological 

genomics—changing perspectives on Darwin’s basic concerns.

Molecular Ecology 19:3025-3030.

Type: Journal 

Geographic Area: Global

Keywords: adaptation, gene duplication, gene expression, 

gene phylogeny, genetic variation, genomics, speciation 

Abstract: Ecological Genomics is an interdisciplinary field 

that seeks to understand the genetic and physiological basis 

of species interactions for evolutionary inferences. At the 

7th annual Ecological Genomics Symposium, November 

13–15, 2009, members of the Ecological Genomics program 

at Kansas State University invited 13 speakers and 56 poster 

presentations.

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CHAPTER 2 • CONSERVATION AND RESTORATION • STRATEGIES

loss of buffering against environmental variation as the dormant seed bank was reduced. Our stochastic simulations suggested that extinction risk for Holocarpha populations would 

be minimized by intermediate levels of ray seed germination. 

Thus, managers should focus on improving the performance 

of aboveground plants before considering actions to stimulate 

germination, since the former will yield a greater increase in 

deterministic population growth and not sacrifice any buffering effect of the seed bank. This case study emphasizes the 

importance of considering dormant seeds and seed banks in 

designing successful restoration and management strategies 

for plant species at risk of extinction.

Link: http://www.elkhornsloughctp.org/uploads/

files/1168483236Satterthwaite%202007%20Seed%20Banks.

pdf

316. Schoettle AW. 2004. Developing proactive management options to sustain bristlecone and limber pine ecosystems 

in the presence of a non-native pathogen. In: Shepperd WD, 

Eskew LG, editors. Silviculture in Special Places: Proceedings 

of the National Silviculture Workshop. Fort Collins (CO): 

USDA Forest Service, Rocky Mountain Research Station. 

Proceedings RMRS-P-34. p 146-155.

Type: Government Document

Geographic Area: Northwestern USA

Compilers’ Keywords: white pine blister rust, strategy, 

Rocky Mountain, Pinus strobus, Pinus flexilis, Pinus aristata

Abstract: Limber pine and Rocky Mountain bristlecone pine 

are currently threatened by the non-native pathogen white pine 

blister rust (WPBR). Limber pine is experiencing mortality in 

the northern Rocky Mountains and the infection front continues to move southward. The first report of WPBR on Rocky 

Mountain bristlecone pine was made in 2003, at a site that is 

more than 220 miles away from the former infection front. 

No mortality has been observed in this recently infected area 

but the species is highly susceptible. There are no ecological 

reasons to suspect that WPBR on bristlecone and the southern 

distribution of limber pine will not expand over time. Learning 

from experiences in impacted ecosystems will facilitate the 

development of proactive measures to mitigate impacts in 

these southern populations in the future. If no action is taken, 

and the pathogen takes its course, we risk losses of aesthetic 

landscapes; impacts to ecosystem boundaries, successional 

pathways, and watershed processes; and shifts from forested 

to treeless sites at some landscape positions. This paper introduces an interdisciplinary approach to developing proactive 

management options for limber and bristlecone pines in the 

southern Rocky Mountains. Managers, researchers, operational professionals and interested public groups will have to work 

together and share their knowledge and perspectives to sustain 

these ecosystems for future generations.

Link: http://www.treesearch.fs.fed.us/pubs/7221

for bioresource development, restoration, and use. As biotechnologies advance, we will appreciate more fully the true costs 

of constructing genetic moieties into fully functional organisms, and maintaining populations of these under controlled 

conditions or combining them in self-perpetuating systems. 

Intact populations and ecosystems will be ever more valuable 

sources of the genetic resources and contextual information 

required for this enterprise. Three elements are required to 

realize the values of biodiversity: genetic materials; environments; and information about the functional relationships of 

the first two. Effective conservation involves uniting these 

elements. Thorough review of historical, scientific, socioeconomic, and practical information about particular elements 

of biodiversity is necessary to assess the constraints on and 

opportunities for conservation activities. The California Gene 

Resources Program activities from 1980 to 1983 are examples 

of this kind of undertaking. Computerized information systems 

can assist managers and researchers in uniting these elements 

and facilitating both conservation of, and access to, genetic 

resources. The California Forest Genetic Sources Catalog, a 

microcomputer database application developed by GENREC 

for the Wildland Resources Center, is one such application. 

Well-constructed databases coupled with knowledge-based 

decision aids will become indispensable as in situ preserves 

and managed areas are integrated with ex-situ collections and 

research findings into effective genetic-resource conservation 

systems.

315. Satterthwaite W, Holl K, Hayes G, Barber A. 

2007. Seed banks in plant conservation: case study of Santa 

Cruz tarplant restoration. Biological Conservation 135:57-66.

Type: Journal 

Geographic Area: California, USA

Keywords: dormancy, population viability, coastal prairie, 

reintroduction macradenia

Abstract: Although conservation and restoration practitioners have focused on maximizing aboveground population 

size and seed set of rare plants, a clear understanding of seed 

bank dynamics is crucial to managing these species. Santa 

Cruz tarplant (Holocarpha macradenia) is a threatened annual 

forb restricted to coastal prairie habitats in central California. 

Holocarpha produces disk achenes germinating within a year 

of production and ray achenes forming a persistent seed bank. 

We constructed both deterministic and stochastic demographic 

models for a restored Holocarpha population, using demographic rates measured separately for unmanipulated plants 

and plants growing in plots where vegetation was clipped. The 

deterministic models indicated that regardless of germination 

from the seed bank, the population would decline without clipping or similar treatments that enhance survival and reproductive output. Deterministic models showed only a slight positive 

effect of increased ray seed germination rates on population 

growth, which would need to be balanced against a potential 
114 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

CHAPTER 2 • CONSERVATION AND RESTORATION • STRATEGIES

upon invasion is also an immediate need—as the disease intensifies, time is running out. 

Link: http://www.treesearch.fs.fed.us/pubs/29501

319. Schoettle AW, Goodrich BA, Klutsch JG, 

Burns KS, Costello S, Sniezko R, Connor J. 2011. 

The proactive strategy for sustaining five-needle pine populations: an example of its implementation in the Southern Rocky 

Mountains. In: Keane RE, Tomback DF, Murray MP, Smith 

CM, editors. High Five Symposium. Fort Collins (CO): USDA 

Forest Service, Rocky Mountain Research Station. Proceedings 

RMRS-P-63. p 323-334.

Type: Government Document

Geographic Area: Northwestern USA

Compilers’ Keywords: Cronartium ribicola, white pine blister rust, mountain pine beetle, Dendroctonus ponderosae, climate change, limber pine, management, genetic diversity, seed 

collection

Abstract: The imminent invasion of the non-native fungus, Cronartium ribicola J.C. Fisch., that causes white pine 

blister rust (WPBR) and the current mountain pine beetle 

(Dendroctonus ponderosae Hopkins, MPB) epidemic in 

northern Colorado limber pine forests will severely affect 

the forest regeneration cycle necessary for functioning 

ecosystems. The slow growth and maturity of limber pine 

enables trees to persist on the landscape for centuries, but 

without sufficient regeneration opportunities these traits 

will inevitably hinder the ability of limber pine to adapt to 

novel stresses such as WPBR or climate change. The current 

MPB outbreak will result in the death of many mature limber 

pines, including many with genetic resistance to WPBR. It 

will be decades until advanced regeneration develops into 

seed-producing mature trees in much of this region. This 

development will be limited further by WPBR which rapidly kills susceptible young trees. Efforts to sustain limber 

pine and Rocky Mountain bristlecone pine in the southern 

Rockies were initiated in 2001. The Northern Colorado 

Limber Pine Conservation Program, described here, is an example of the Proactive Strategy being implemented at a local 

scale. The program is a cooperative partnership between the 

USFS Rocky Mountain Research Station, Rocky Mountain 

National Park (RMNP), USFS Forest Health Management, 

and several Ranger Districts on the Arapaho-Roosevelt 

National Forest. It is designed to 1) conserve limber pine 

genetic diversity and 2) collect needed data to assist land 

managers in making informed decisions for preparation of 

a management plan intended to sustain resilient limber pine 

ecosystems in northern Colorado. The cooperative effort 

aims to provide immediate protection of limber pine from 

MPB, facilitate seed collections for WPBR resistance tests, 

and store and use seed for conservation, research and restoration. Seedlings are being screened for WPBR resistance 

317. Schoettle AW, Sniezko RA. 2007. Proactive intervention to sustain high-elevation pine ecosystems threatened by 

white pine blister rust. Journal of Forest Research 12:327-336.

Type: Journal 

Geographic Area: Western North America

Keywords: evolution of resistance, exotic pathogen, Pinus 

aristata, Pinus albicaulis, Pinus flexilis

Abstract: Only recently have efforts begun to address how 

management might prepare currently healthy forests to affect the outcome of invasion by established nonnative pests. 

Cronartium ribicola, the fungus that causes the disease white 

pine blister rust (WPBR), is among the introductions into 

North America where containment and eradication have 

failed; the disease continues to spread. Ecosystem function 

is impaired by high rust-caused mortality in mature fiveneedle white pine forests. This paper evaluates five proactive management options to mitigate the development of impacts caused by white pine blister rust in threatened remote 

high-elevation five-needle pine ecosystems of western North 

America. They are: reducing pest populations; managing forest composition; improving host vigor; introducing resistant 

stock with artificial regeneration; and diversifying age class 

structure to affect the natural selection process for resistance. 

Proactive intervention to manage and facilitate evolutionary 

change in the host species may sustain host populations and 

ecosystem function during pathogen naturalization.

Link: http://www.treesearch.fs.fed.us/pubs/29500

318. Schoettle AW, Sniezko R, Burns KS, Freeman 

F. 2007. Preparing the landscape for invasion—early intervention approaches for threatened high elevation white pine ecosystems. In: Goheen EM, Sniezko RA, editors. Whitebark Pine: 

A Pacific Coast Perspective. Portland (OR): USDA Forest 

Service, Pacific Northwest Region. R6-NR-FHP-2007-01. p 

72-75.

Type: Government Document

Geographic Area: Northwestern USA

Compilers’ Keywords: whitebark pine, Pinus albicaulis, 

white pine blister rust, regeneration, forest management, pest 

management

Introduction: White pine blister rust is now a permanent 

resident of North America. The disease continued to cause tree 

mortality and impact ecosystems in many areas. However, not 

all high elevation white pine ecosystems have been invaded; 

the pathogen is still spreading within the distributions of the 

whitebark, limber, foxtail, Rocky Mountain bristlecone pine 

and has yet to infect Great Basin bristlecone pines. While the 

heavily impacted areas are in need of immediate management 

to restore ecosystem function, management of the threatened 

areas to position them to avoid development of severe impacts 
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 115

CHAPTER 2 • CONSERVATION AND RESTORATION • STRATEGIES

Link: http://www.treesearch.fs.fed.us/pubs/39211

321. Shaw NL, Lambert SM, DeBolt AM, Pellant 

M. 2005. Increasing native forb seed supplies for the Great 

Basin. In: Dumroese RK, Riley LE, Landis TD, editors. Forest 

and Conservation Nursery Associations—2004. Fort Collins 

(CO): USDA Forest Service, Rocky Mountain Research 

Station. Proceedings RMRS-P-35. p 94-102.

Type: Government Document

Geographic Area: Great Basin USA

Keywords: native forbs, plant materials, seed production, 

seeding, rangelands

Abstract: Over the last 150 years, excessive grazing, annual 

weed invasions, increased wildfire frequency, and other human 

disturbances have negatively impacted native plant communities of the Great Basin. Native plant materials and appropriate 

planting strategies are needed to recreate diverse communities 

in areas requiring active restoration. Although native forbs 

are critical components of most plant communities, available 

seed supplies remain low. A cooperative research project being 

conducted by the USDI Bureau of Land Management Great 

Basin Restoration Initiative, the USDA Forest Service Rocky 

Mountain Research Station, and collaborators includes efforts 

to develop 20 native forbs as revegetation species. Research 

needs include selection of seed sources and development of 

seed production and wildland seeding technology for each species. Initial seed increase of new seed sources and maintenance 

of seed supplies will require production at a range of scales, 

likely creating new marketing niches for the native seed and 

nursery industries.

Link: http://www.treesearch.fs.fed.us/pubs/20897

322. St. Clair JB, Lipow S, Vance-Borland K, Johnson 

R. 2007. Conservation of forest genetic resources in the United 

States. In: Loo JA, Simpson JD, editors. Proceedings of the 

13th Meeting of the Canadian Tree Improvement Association: 

Canada’s Forests-Enhancing Productivity, Protection and 

Conservation. New Brunswick, Canada. p 16-24.

Type: Conference Paper

Geographic Area: USA

Compilers’ Keywords: genetic variation, forest management, gene conservation, gap analysis, tree improvement

Abstract: Conservation of genetic diversity is recognized 

as an important requirement of sustainable forest management. Gene conservation activities include in situ conservation of native stands in reserves and ex situ conservation in 

seed banks, genetic tests, seed and breeding orchards, and 

other plantations of known identity. We present an example 

from Oregon and Washington of a GIS-based “gap analysis” 

approach to determine the spatial distribution of protected 

in situ genetic resources. GIS coverages showing detailed 

to determine frequencies of resistance across the landscape 

among populations and to identify resistant seed trees for 

future seed collections. The seedling tests will also estimate 

population differentiation along the elevation gradient to refine seed transfer guidelines. Surveys of forest health, biotic 

damage, rust incidence, and trends in advanced regeneration 

will help project persistence of these populations after passage of the MPB epidemic. These data will provide a basis for evaluations of proactive management options on a 

site specific basis before northern Colorado ecosystems are 

impaired by WPBR; this should shorten the time frame to 

return to functioning ecosystems. Focusing management on 

proactively maintaining genetic diversity and a functional 

regeneration cycle will promote sustained adaptive capacity 

and ecosystems resilience to novel stresses such as WPBR 

and climate change.

Link: http://www.treesearch.fs.fed.us/pubs/38244

320. Shaw NL, Youtie B, Olwell P. 2011. Building 

bridges between agencies, researchers, farmers and non-governmental organizations to create collaborative native seed 

programs. In: Proceedings 7th European Conference on 

Ecological Restoration. Avignon, France: Society for 

Ecological Restoration. p 1-4.

Type: Conference Paper

Geographic Area: Western USA

Keywords: climate change, disturbance by fire, invasive species, native plant programs

Abstract: The Native Plant Materials Development Program 

was authorized by the U.S. Department of the Interior and 

Related Agencies Appropriations Act of FY2001 to provide support for development of native plant materials required for restoration of disturbed public lands in the U.S.A. 

The Washington, DC, Office of the USDI Bureau of Land 

Management has provided national leadership for this program 

by developing partnerships with more than 250 public and private entities, a national program for collecting germplasm for 

ex situ conservation and for seed increase for restoration uses, 

and by initiating state and regional level native plant materials 

programs. Regional programs such as the Great Basin Native 

Plant Selection and Increase Project aim to fulfill long-term 

plant material needs by providing genetically diverse, regionally appropriate seed sources and fostering the research 

required to produce seeds of new restoration species in agricultural settings. The Deschutes Basin Native Plant Seedbank, 

a non-profit organization, has provided valuable leadership by 

functioning as a buyer’s cooperatives to provide affordable local seed for public and private partners in areas with a mosaic 

of land ownerships. At all levels, native plant materials development programs rely heavily upon cooperation among land 

managers, researchers, non-governmental organizations, and 

the private sector seed industry.
116 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

CHAPTER 2 • CONSERVATION AND RESTORATION • STRATEGIES

1 to 2 wk after burning reduced broadleaf weeds, and seed 

addition increased native diversity.

Link: http://www.researchgate.net/publication/228743458_

Regional_strategies_for_restoring_invaded_prairies_observations_from_a_multi-site_collaborative_research_project/

file/9fcfd5059e6ca4a2cd.pdf

324. Thomas E, Jalonen R, Loo J, Boshier D, Gallo 

L, Cavers S, Bordács S, Smith P, Bozzano M. 2014.

Genetic considerations in ecosystem restoration using native tree 

species. Forest Ecology and Management 333:66-75.

Type: Journal 

Geographic Area: Global

Keywords: ecosystem restoration, genetic diversity, native 

tree species, seed selection, measures of success, tree nursery

Abstract: Rehabilitation and restoration of forest ecosystems are in growing demand to tackle climate change, 

biodiversity loss and desertification—major environmental 

problems of our time. Interest in restoration of ecosystems 

is increasingly translated into strong political commitment 

to large-scale tree planting projects. Along with this new 

impetus and the enormous scale of planned projects come 

both opportunities and risks: opportunities to significantly 

increase the use of native species, and risks of failure associated with the use of inadequate or mismatched reproductive material, which though it may provide forest cover 

in the short term, will not likely establish a self-sustaining 

ecosystem. The value of using native tree species in ecosystem restoration is receiving growing recognition both among 

restoration practitioners and policy makers. However, insufficient attention has been given to genetic variation within 

and among native tree species, their life histories and the 

consequences of their interactions with each other and with 

their environment. Also restoration practitioners have often 

neglected to build in safeguards against the anticipated effects of anthropogenic climate change. Measurement of 

restoration success has tended to be assessments of hectares 

covered or seedling survival in a short timeframe, neither of 

which is an indicator of ecosystem establishment in the long 

term. In this article, we review current practices in ecosystem restoration using native tree species, with a particular 

focus on genetic considerations. Our discussion is organised 

across three themes: (i) species selection and the sourcing of 

forest reproductive material; (ii) increasing resilience by fostering natural selection, ecological connectivity and species 

associations; and (iii) measuring the success of restoration 

activities. We present a number of practical recommendations for researchers, policymakers and restoration practitioners to increase the potential for successful interventions. 

We recommend the development and adoption of decisionsupport tools for: (i) collecting and propagating germplasm 

in a way that ensures a broad genetic base of restored tree 

distributions of eight tree species stratified into presumably 

unique genetic units using seed zones or ecoregions were 

overlaid with coverages of reserves to determine the locations of protected populations as Well as gaps in protection. 

The gap analysis indicated that most species appear to have 

sufficient genetic resources conserved in in situ reserves. 

This approach may be valuable elsewhere, particularly in 

the eastern United States where fewer large reserves exist. Of particular note, for ex situ conservation, is a recent 

agreement between the USDA Forest Service National Seed 

Laboratory and the USDA Agricultural Research Service 

National Center for Genetic Resources Preservation to 

provide long-term storage of seed collections of valuable 

plant germplasm in their facility in Ft. Collins, Colorado. 

Protection from disease and insects is another important 

component of gene conservation.

Link: http://www.treesearch.fs.fed.us/pubs/29348

323. Stanley AG, Kaye TN, Dunwiddle PW. 2008.

Regional strategies for restoring invaded prairies: observations 

from a multisite, collaborative research project. Native Plants 

Journal 9:247-254.

Type: Journal 

Geographic Area: Pacific Northwest USA

Keywords: grassland restoration, prairie, herbicide, restoration methods, seed addition, native diversity, invasive species

Abstract: Invasive plants, especially nonnative perennial 

grasses, pose one of the most critical threats to protected 

prairies and oak woodlands in the Pacific Northwest. Our 

current knowledge regarding the effectiveness of weed 

control methods, especially in sites that retain a significant 

component of native vegetation, is largely anecdotal or 

based on results from a few site-specific studies. The Nature 

Conservancy jointly with the Institute for Applied Ecology 

and its partners have initiated a large-scale, long-term, interdisciplinary, and collaborative project to: 1) evaluate and 

improve strategies for controlling the abundance of invasive 

nonnative herbaceous weeds while maintaining or enhancing the abundance and diversity of native plant species; and 

2) develop an approach to generalize results so they can be 

applied by land managers engaged in prairie stewardship 

throughout the region. This project combines simultaneous 

small-scale replicated experiments with large-scale unreplicated experiments at 11 sites in Washington, Oregon, and 

British Columbia. Experimental treatments, begun in 2005, 

include combinations of spring and fall mowing, burning, 

a grass-specific herbicide (sethoxydim), a broad-spectrum 

herbicide (glyphosate), and seeding of native species. Our 

preliminary observations show sethoxydim applications 

effectively reduce exotic perennial grasses. Combining sethoxydim with other treatments had added benefits: fall burning reduced thatch and moss cover, glyphosate application 
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 117

CHAPTER 2 • CONSERVATION AND RESTORATION • STRATEGIES

populations, including planning the sourcing of propagation 

material of desired species well before the intended planting 

time; (ii) matching species and provenances to restoration 

sites based on current and future site conditions, predicted 

or known patterns of variation in adaptive traits and availability of seed sources; and (iii) landscape-level planning in 

restoration projects.

Link: http://www.sciencedirect.com/science/article/pii/

S0378112714004356

325. van Slageren W. 2003. The Millennium Seed Bank: 

building partnerships in arid regions for the conservation of wild 

species. Journal of Arid Environments 54:195-201.

Type: Journal 

Geographic Area: United Kingdom, Global

Keywords: seed bank, conservation, ex-situ, drylands, benefit-sharing, conventions, research, education

Abstract: The Millennium Seed Bank Project is a large, 

international conservation project. Most of the project will 

focus—in collaboration with many dryland countries—on 

the much-neglected need for conservation of wild species in 

the (semi-) arid regions of the world. Its principal aim is to 

help safeguard 24,000 species of dryland plants—10% of the 

world’s flora—against extinction. A second aim is, equally 

through ex-situ conservation, to secure the future of almost 

all of the U.K.’s native flowering plants. For the project, a 

new building has been constructed at Wakehurst Place in 

Sussex, part of the Royal Botanic Gardens, Kew. After opening in late 2000, it will house the Seed Bank and be a world 

resource for seed conservation, research and education.

Link: http://www.cbd.int/doc/articles/2003/A-00155.pdf

326. Zhou S, Sauve RJ. 2006. Genetic fingerprinting of 

goldenseal using AFLP markers. Native Plants Journal 7:73-77.

Type: Journal 

Geographic Area: Southeastern USA

Keywords: genetic diversity, molecular marker, plant DNA 

profile

Abstract: Goldenseal (Hydrastis canadensis L. 

[Ranunculaceae]) could be considered a worthy ornamental 

plant for a forest-like garden setting, however, its primary 

use is medicinal. Research described in this paper demonstrates that AFLP analysis can be used to determine the 

genetic relationships between accessions of a plant species 

collected from different regions. The genetic relationships 

of 9 goldenseal accessions collected in 3 neighboring states, 

Tennessee, Georgia, and Florida, were determined. A cluster analysis from AFLP data showed that the 2 Tennessee 

accessions were very closely related to each other with a 

high bootstrap value of 93%. The 4 Georgia accessions were 

more diversified with a bootstrap value ranging from 38% to 

63%. Among the 3 Florida accessions, genetic relatedness 

was very low. One Florida accession was closely related to 

the 2 Tennessee accessions (96%), one with the Georgia accessions (88%), and the other was distant to all accessions. 

The molecular marker technique developed for determining 

the genetic relationship and the genetic diversity between 

accessions could be used by plant breeders for the selection 

of parental material.
118 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

CHAPTER 2 • CONSERVATION AND RESTORATION • RESOURCES

327. Bonner FT, Karrfalt RP. 2008. The woody plant 

seed manual. Washington (DC): USDA Forest Service. 

Agriculture Handbook 727. 1223 p.

Type: Government Document

Geographic Area: USA

Compilers’ Keywords: seed storage, seed collection, germination, handbook, propagation, outplanting

Introduction: The first comprehensive handbook on the 

seeds of trees and shrubs produced by the USDA Forest Service 

was USDA Misc. Pub. 654, Woody-Plant Seed Manual. The 

manuscript was ready for publication in 1941, but World War 

II delayed publication until 1948. The boom in tree planting 

in the 1950s and 1960s created a large demand for seeds and 

exposed the gaps in our knowledge concerning production and 

quality of seeds of woody plants in general. Realization of this 

condition led to the revision and considerable expansion of 

the manual, resulting in publication of USDA Agric. Handbk. 

450, Seeds of Woody Plants in the United States, in 1974. Seed 

data were presented for about 800 species, varieties, and subspecies in 188 genera, considerable more than the 420 species 

and 140 genera in the 1948 edition. The 1974 Handbook proved 

to be very popular both in this country and abroad, leading to 

five printings and translations in several other languages. More 

than a quarter-century after its publication, however, numerous 

advances in tree seed technology have dictated that a new revision is needed; the result is the current volume.

Link: http://www.treesearch.fs.fed.us/pubs/32626

328. Dumroese RK, Owston P. 2003. A user’s guide to 

nursery stock types. Western Forester 4-5.

Type: Newsletter

Geographic Area: USA

Compilers’ Keywords: nursery practices, user guide, forest 

management, seedlings, trees, container

Introduction: Foresters must consider many factors when 

selecting nursery stock types for their planting projects. 

Forest nurseries can now produce a vast array of stock types 

to meet any challenge in the field—these target seedlings 

can be defined for particular sites. It is important that foresters work with a nursery that is known from experience 

or reputation to provide a consistently good product at a 

reasonable price. Foresters and nurseries must foster open 

and effective communication to continually improve stock 

type quality. To facilitate communication between nurseries 

and foresters, a system of naming stock types developed and 

General (215–232)—reviews, surveys, 

summaries, theory, foundations, paradigms

Research (233–285)—propagation, 

outplanting, invasion

Strategies (286–326)—frameworks, approach, 

decision support

Resources (327–338)—native plant materials, 

guidelines, handbooks

Conservation and 

Restoration

#. Author(s), Date. Title. Source.

Type: 

Geographic Area: 

Keywords: 

Description:

Link:

location of study

keywords written by the 

author(s) or provided by the 

compilers

ID number and

information

abstract, summary, or 

portions of the introduction 

or conclusion written by the 

author(s) or provided by the 

compilers

book, book section, conference 

paper, dissertation, government 

document, journal article, 

newsletter, presentation, report, 

thesis or web page

internet address of reference

Each reference contains the following:
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 119

project with Spain to promote the training and research of plant 

genetic resources in Latin America. The text trains users in the 

fundamental aspects of ex situ conservation of plant genetic 

resources, that is, from collecting to germplasm use. It explains 

principles and describes the procedures needed for effective ex 

situ conservation. It also includes bibliographic references and 

examples that illustrate how to move from theory to practice. 

By developing and making this material available to users, 

Biodiversity International hopes to significantly contribute to 

the training of technicians in plant genetic resources, thereby 

making them more efficient in ex situ conservation and germplasm use.

331. McCaughey W, Scott GL, Izlar KL. 2009.

Whitebark pine planting guidelines. Western Journal of Applied 

Forestry 24:163-166.

Type: Journal 

Geographic Area: Northern USA

Keywords: Pinus albicaulis, reforestation, tree-planting, seedlings, plantations

Abstract: This article incorporates new information into 

previous whitebark pine guidelines for planting prescriptions. 

Earlier 2006 guidelines were developed based on review of 

general literature, research studies, field observations, and 

standard U.S. Forest Service survival surveys of high-elevation whitebark pine plantations. A recent study of biotic and 

abiotic factors affecting survival in whitebark pine plantations 

was conducted to determine survival rates over time and over a 

wide range of geographic locations. In these revised guidelines, 

we recommend reducing or avoiding overstory and understory 

competition, avoiding swales or frost pockets, providing shade 

and wind protection, protecting seedlings from heavy snow 

loads and soil movement, providing adequate growing space, 

avoiding sites with lodgepole or mixing with other tree species, and avoiding planting next to snags.

Link: http://www.fs.fed.us/rm/pubs_other/rmrs_2009_mccaughey_w001.pdf

332. Monsen SB, Stevens R, Shaw NL. 2004.Grasses.

Chapter 18. In: Monsen SB, Stevens R, Shaw NL, editors. 

Restoring Western Ranges and Wildlands. Fort Collins (CO): 

USDA Forest Service, Rocky Mountain Research Station. 

General Technical Report RMRS-GTR-136. p 295-424. 

Type: Government Document

Geographic Area: Western USA

Compilers’ Keywords: plant ecology, native plant material, 

wildlife habitat, restoration

Introduction: Grasses are adapted to a wide range of edaphic 

and climatic conditions and are found in nearly all plant communities. In the Western United States, grasses are seeded on 

disturbances to provide forage, wildlife habitat, and watershed 

those stock type descriptions fall into three broad categories: 

bareroot, container and “plug plus.”

Link: http://www.treesearch.fs.fed.us/pubs/32849

329. Dumroese RK, Landis TD, Luna T. 2012.

Raising native plants in nurseries: basic concepts. Fort Collins 

(CO): USDA Forest Service Rocky Mountain Research 

Station. General Technical Report RMRS-GTR-274. 92 p.

Type: Government Document

Geographic Area: North America

Compilers’ Keywords: nursery practices, propagation, greenhouse, seed, container nursery, bareroot nursery, guidelines

Abstract: Growing native plants can be fun, challenging, and 

rewarding. This booklet, particularly the first chapter that introduces important concepts, is for the novice who wants to 

start growing native plants as a hobby; however, it can also be 

helpful to someone with a bit more experience who is wondering about starting a nursery. The second chapter provides basic 

information about collecting, processing, storing, and treating 

seeds. Chapter three focuses on using seeds to grow plants in 

the field or in containers using simple but effective techniques. 

For those native plants that reproduce poorly from seeds, the 

fourth chapter describes how to start native plants from cuttings. The final chapter provides valuable information on how 

to successfully move native plants from the nursery and establish them in their final planting location. Several appendices 

expand on what has been presented in the chapters, with more 

details and specific information about growing a variety of native plants.

Link: http://www.treesearch.fs.fed.us/pubs/40753

330. Jaramillo S, Baena M. 2007. Ex situ conservation 

of plant genetic resources. Cali, Columbia: International Plant 

Genetic Resources Institute. 229 p. 

Type: Book

Geographic Area: Global

Compilers’ Keywords: Latin America, Spain, training module, guide

Forward: Plant genetic resources form the basis on which 

humankind subsists. They provide basic needs and help solve 

problems such as hunger and poverty. However, they are being lost, mainly through inappropriate use and destruction of 

habitat. Given their vital importance, we must conserve them 

for the benefit of both present and future generations. Plant 

genetic resources can be conserved within or outside their 

natural habitats, or by combining the two alternatives. Outside 

their natural habitats, plant genetic resources are conserved in 

germplasm collections and genebanks, going through different 

stages and procedures that require trained staff. To help train 

personnel in managing genebanks, Bioversity International 

has developed this teaching manual as part of a collaborative 

CHAPTER 2 • CONSERVATION AND RESTORATION • RESOURCES
120 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

native populations of Lewis flax. All available data indicate 

that ‘Appar’ is not invasive and does not pose a threat to native 

flax populations. Ease of establishment, seed production, and 

showiness of the abundant flowers recommend ‘Appar’ for 

use in xeriscape plantings. Alternatively, the recent germplasm 

release of Lewis flax, ‘Maple Grove’, also establishes readily 

from seed and can be effectively used when objectives dictate 

the exclusive use of native germplasms.

Link: http://www.treesearch.fs.fed.us/pubs/30681

335. Proctor J, Haas W. 2007. Guidelines for revegetation for the Medicine Bow-Routt National Forests and Thunder 

Basin National Grassland. 69 p.

Type: Report

Geographic Area: Intermountain West USA

Compilers’ Keywords: restoration, handbook

Compilers’ Summary: This is a revegetation handbook for 

the Medicine Bow-Routt National Forests and Thunder Basin 

National Grassland that cover approximately 2.9 million 

acres in north central Colorado and central and northeastern 

Wyoming. It covers guidelines for mountain ranges, grasslands and riparian systems. 

336. Scott GL, McCaughey WW. 2006.Whitebark pine 

guidelines for planting prescriptions. In: Riley LE, Dumroese 

RK, Landis TD, editors. National Proceedings: Forest and 

Conservation Nursery Associations—2005. Fort Collins (CO): 

USDA Forest Service, Rocky Mountain Research Station. 

Proceedings RMRS-P-43. p 84-90.

Type: Government Document

Geographic Area: Western USA

Keywords: whitebark pine, Pinus albicaulis, reforestation, tree 

planting, pocket gophers, white pine blister rust, mountain pine 

beetle, cones, seeds, fire suppression

Abstract: This paper reviews general literature, research 

studies, field observations, and standard Forest Service survival surveys of high-elevation whitebark pine plantations and 

presents a set of guidelines for outplanting prescriptions. When 

planting whitebark pine, the recommendations are: 1) reduce 

overstory competition; 2) reduce understory vegetation, especially grasses and sedges; 3) avoid outplanting in swales or 

frost pocket areas; 4) provide shade protection; 5) plant where 

there is protection from heavy snow loading; and 6) provide 

adequate growing space. 

Link: http://www.treesearch.fs.fed.us/pubs/26661

337. Society for Ecological Restoration Ecological 

Restoration International Science and Policy 

Working Group. 2004. The SER international primer on 

ecological restoration. Tucson (AZ): Society for Ecological 

Restoration International. 15 p.

stability. A number of introduced grass species proved wellsuited to Western rangelands and received extensive use in 

early reseeding efforts. Use of these and other introductions 

continued over time, but by the late 1900s, greater emphasis 

was being placed on the use of native grasses. 

Link: http://www.treesearch.fs.fed.us/pubs/31963

333. Monsen SB, Stevens R, Shaw NL. 2004.

Restoring western ranges and wildlands. Fort Collins (CO): 

USDA Forest Service, Rocky Mountain Research Station. 

General Technical Report RMRS-GTR-136. p 699-884.

Type: Government Document

Geographic Area: Western USA

Keywords: rehabilitation, revegetation, plant ecology, seed, 

plant communities, wildlife habitat, invasive species, equipment, plant materials, native plants

Abstract: This work, in three volumes, provides background 

on philosophy, processes, plant materials selection, site preparation, and seed and seeding equipment for revegetating disturbed rangelands, emphasizing use of native species. The 

29 chapters include guidelines for planning, conducting, and 

managing, and contain a compilation of rangeland revegetation research conducted over the last several decades to aid 

practitioners in reestablishing healthy communities and curbing the spread of invasive species. Volume 3 contains chapters 

24-29 plus appendices and index.

Link: http://www.treesearch.fs.fed.us/pubs/7379

334. Pendleton RL, Kitchen SG, McArthur ED, 

Mudge JE. 2008. The ‘Appar’ flax release: origin, distinguishing characteristics, and use; and a native alternative.

Native Plants Journal 9:18-24.

Type: Journal 

Geographic Area: North America, Europe

Keywords: heterostyly, distyly, native plant material, revegetation, xeriscaping, Linum lewisii, Linum perenne, Linaceae, 

native germplasm

Abstract: This article summarizes information on the taxonomy of ‘Appar’, a perennial blue flax cultivar (Linum perenne

L. [Linaceae]), and characteristics that distinguish it from 

native Lewis flax (Linum lewisii Pursh [Linaceae]). ‘Appar’ 

apparently originated as a European flax that escaped from 

garden cultivation. Randomly amplified polymorphic DNA 

(RAPD) analysis places ‘Appar’ with other collections of 

European Linum perenne and separates it from native North 

American collections of Lewis flax. Morphologically, ‘Appar’ 

differs from Lewis flax in having an intense blue petal color, 

shorter internodes, leaves that point upward along the stem, 

and multiple slender stems suffused with red near the base. 

The heterostylous reproductive system, which it shares with its 

European cousins, keeps ‘Appar’ reproductively isolated from 

CHAPTER 2 • CONSERVATION AND RESTORATION • RESOURCES
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 121

CHAPTER 2 • CONSERVATION AND RESTORATION • RESOURCES

Type: Report

Geographic Area: Global

Compilers’ Keywords: planning, manual, guidelines

Introduction: Ecological restoration is an intentional activity 

that initiates or accelerates the recovery of an ecosystem with 

respect to its health, integrity and sustainability. Frequently, the 

ecosystem that requires restoration has been degraded, damaged, transformed or entirely destroyed as the direct or indirect 

result of human activities. In some cases, these impacts to ecosystems have been caused or aggravated by natural agencies 

such as wildfire, floods, storms, or volcanic eruption, to the 

point at which the ecosystem cannot recover its predisturbance 

state or its historic developmental trajectory. This document is 

a primer on ecological restoration.

Link: http://www.ser.org/resources/resources-detail-view/serinternational-primer-on-ecological-restoration

338. USDA RNGR. 2014. National Nursery and Seed 

Directory. (URL accessed 29 November 2015)

Type: Web Page 

Geographic Area: USA

Compilers’ Keywords: native plants, directory, plant 

materials

Summary: This is a directory of nurseries and farms that produce native plant materials. It is maintained by the USDA Forest 

Service, Reforestation, Nurseries, and Genetics Resources 

(RNGR) team in collaboration with the University of Georgia, 

Southern Regional Extension Forestry (SREF). SREF verifies 

one-third of the directory annually. The RNGR website (http://

www.rngr.net), also holds a vast amount of information about 

growing native plants. One of the most important differences 

between native plant materials and ornamental nursery stock is 

that natives are usually outplanted in relatively harsh environments generally without any subsequent care. 

Link: http://www.rngr.net/resources/directory
122 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

CHAPTER 3 • MIGRATION • GENERAL

339. Albrecht GA, Brooke C, Bennett DH, Garnett 

ST. 2012. The ethics of assisted colonization in the age of 

anthropogenic climate change. Journal of Agricultural and 

Environmental Ethics 26:827-845.

Type: Journal 

Geographic Area: Global

Keywords: ethics, climate change, assisted colonization, managed relocation, mitigation, adaptation, intrinsic value, instrumental value, sentience, environmental ethics, sustainability 

ethics, review

Abstract: This paper examines an issue that is becoming increasingly relevant as the pressures of a warming planet, changing climate and changing ecosystems ramp up. The broad context for the paper is the intragenerational, intergenerational, and 

interspecies equity implications of changing the climate and the 

value orientations of adapting to such change. In addition, the 

need to stabilize the planetary climate by urgent mitigation of 

change factors is a foundational ethical assumption. In order to 

avoid further animal and plant extinctions, or at the very least, 

their increased vulnerability to becoming rare and endangered; 

the systematic assisted colonization of “at risk” species is being 

seriously considered by scientists and managers of biodiversity. 

The more practical aspects of assisted colonization have been 

covered in the conservation biology literature; however, the 

ethical implications of such actions have not been extensively 

examined. Our discussion of the value issues, using a novel case 

study approach, will rectify the limited ethical analysis of the 

issue of assisted colonization of species in the face of climate 

change pressures. 

340. Aubin I, Garbe CM, Colombo S, Drever CR, 

McKenney DW, Messier C, Pedlar J, Sander MA, 

Venier L, Wellstead AM, Winder R, Witten E, 

Ste-Marie C. 2011. Why we disagree about assisted migration: ethical implications of a key debate regarding the future of 

Canada’s forests. The Forestry Chronicle 87:755-765.

Type: Journal 

Geographic Area: Canada

Keywords: assisted colonization, climate change, environmental ethics, forest management, species conservation

Abstract: Assisted migration has been proposed as one tool 

to reduce some of the negative ecological consequences of 

climate change. The idea is to move species to locations that 

could better suit them climatically in the future. Although human mediated movements are not a recent phenomenon, assisted migration has lately been the source of debate, in particular within conservation biology circles. In this paper, we 

General (339–380)—debates, reviews, 

trends, surveys, summaries

Research (381–540)—migration studies, 

range shifts, reintroductions

Strategies (541–580)—adaptation options, 

frameworks, decision support

Resources (581–582)—tools, websites, 

software

Migration

#. Author(s), Date. Title. Source.

Type: 

Geographic Area: 

Keywords: 

Description:

Link:

location of study

keywords written by the 

author(s) or provided by the 

compilers

ID number and

information

abstract, summary, or 

portions of the introduction 

or conclusion written by the 

author(s) or provided by the 

compilers

book, book section, conference 

paper, dissertation, government 

document, journal article, 

newsletter, presentation, report, 

thesis or web page

internet address of reference

Each reference contains the following:
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 123

CHAPTER 3 • MIGRATION • GENERAL

is also plausible to posit that the reason this particular species 

failed to make the return journey north is that, in contrast to 

previous interglacial episodes, newly arrived humans obstructed northward migration by overhunting the conifer’s obligate 

seed dispersal partners (squirrels and possibly now-extinct tortoises) in the narrow riverine corridor that contains soils rich 

enough to support the maturation of this yew-like tree.

Link: http://www.torreyaguardians.org/assisted-migration.

html

342. Camacho AE. 2010. Assisted migration: redefining 

nature and natural resource law under climate change. Irvine 

(CA): University of California School of Law. Legal Studies 

Research Paper Series No. 2009-37. 86 p.

Type: Report

Geographic Area: USA

Compilers’ Keywords: land management, resource management, uncertainty, endangered species, economic value conservation, ethics, review

Summary: To avoid extinctions and other harms to ecological health from escalating climatic change, scientists, resource 

managers, and activists are considering and even engaging 

in “assisted migration” - the intentional movement of an organism to an area in which its species has never existed. This 

article explores the profound implications of climate change 

for American natural resource management through the lens of 

this controversial adaptation strategy. It details arguments regarding the scientific viability and legality of assisted migration 

under the thicket of laws that govern natural resources in the 

United States. The article asserts, however, that the fundamental tensions raised by this strategy are ethical: to protect endangered species or conserve native biota; to manage ecological 

systems actively or leave nature wild and uncontrolled; and to 

preserve resources or manage them to promote their fitness under future conditions. The article explains why contemporary 

natural resource law’s fidelity to historic baselines, protecting 

preexisting biota, and shielding nature from human activity is 

increasingly untenable, particularly in light of climate change. 

Active, anticipatory strategies such as assisted migration may 

not only be permissible but even necessary to avert substantial irreversible harm to ecological systems. Scientists and 

resource managers should focus on developing scientific data 

to aid analyses of the risks and benefits of assisted migration 

in particular circumstances. To help develop such data while 

minimizing ecological harm, the article proposes provisionally 

limiting experimental translocations to situations where translocation is technically and economically feasible, and where 

the species is endangered, ecologically valuable, and compatible with the proposed site. More broadly, assisted migration 

illustrates how the institutions and goals of natural resource 

law must be changed to better reflect a dynamic, integrated 

world. Climate changes forces a radical reconsideration of 

outline the major perspectives that help define differing views 

on assisted migration and shed some light on the ethical roots 

of the debate in the context of Canadian forests. We emphasize that there are many different forms of assisted migration, 

each responding to different (often unstated) objectives and 

involving unique risks and benefits, thus making the debate 

more nuanced than often portrayed. We point out certain seeming contradictions whereby the same argument may be used to 

both support and oppose assisted migration. The current debate 

on assisted migration primarily focuses on ecological risks and 

benefits; however, numerous uncertainties reduce our capacity to quantitatively assess these outcomes. In fact, much of 

the debate can be traced back to fundamental perspectives on 

nature, particularly to the ethical question of whether to deliberately manage natural systems or allow them to adapt on their 

own. To facilitate discussion, we suggest that the focus should 

move towards a clearer identification of values and objectives 

for assisted migration.

341. Barlow C. 2011. Paleoecology and the assisted migration debate: why a deep-time perspective is vital. (URL accessed 

15 May 2015) 

Type: Web Page

Geographic Area: Florida, USA

Compilers’ Keywords: Torreya taxifolia, climate change, 

species extinctions

Background and Abstract: In the six years since the assisted migration (assisted colonization) debate was launched 

as a pro and con “Forum” in the Winter 2004/2005 issue of 

Wild Earth journal, the paleoecological perspective has been 

almost entirely missing from both the academic papers and 

the media reports on this vital conservation issue. The original 

PRO-assistance essay by Connie Barlow and (the late) Paul 

S. Martin was titled “Bring Torreya taxifolia North—Now”. 

The original ANTI-assistance essay by Mark Schwartz was 

titled “Conservationists Should Not Move Torreya taxifolia”. 

The intent of this skeletal essay is to demonstrate, by using the 

example of “poster plant” Torreya taxifolia, why the assisted 

migration debate must broaden to encompass a deep-time, paleoecological perspective. Since the beginning of the debate,

the major advocates in favor of assisted migration for Torreya 

taxifolia have argued that paleoecological reasoning is exceedingly strong in favor of assisting the migration of this highly 

endangered conifer tree northward from the panhandle of 

Florida to the southern Appalachians. Notably, there is no dispute that the highly restricted endemic range of Torreya taxifolia in Florida (along the east shore of the Apalachicola River) 

served as one of the primary “pocket refuges” for America’s 

rich Appalachian flora during the peak of the last glacial advance c. 18,000 years ago. It is therefore reasonable to suggest 

that Torreya taxifolia be looked upon as a “glacial relict” that, 

for one reason or another, failed to make the return trip north 

when the present interglacial began some 13,000 years ago. It 
124 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

CHAPTER 3 • MIGRATION • GENERAL

Keywords: adaptation, adaptive peaks, evolution, 

maladaptation

Abstract: This review contains a description of a research 

program for the study of maladaptation, defined here in terms 

of deviation from adaptive peaks. Maladaptation has many genetic causes, including mutation, inbreeding, drift, gene flow, 

heterozygote advantage and pleiotropy. Degrees of maladaptation are determined by genetic architecture and the relationship 

between the rates of selective, environmental change and the 

nature and extent of genetic responses to selection. The empirical analysis of maladaptation requires: (1) recognition of putative maladaptation, using methods from phylogenetics, teleonomy, development and genetics, followed by an assessment 

of the nature and degree of deviation from adaptation, using 

studies of natural selection and teleonomy; (2) determination 

of the causes of the deviation, using analyses of genetics, development, or other methods. Conditions for unambiguously 

identifying maladaptation are considerably more stringent than 

those for demonstrating adaptation and remarkably few studies 

have clearly identified and characterized maladaptive traits. A 

thorough understanding of the nature of phenotypic variation 

will never be achieved without an analysis of the scope and 

usual causes of maladaptation.

Link: http://www.sfu.ca/biology/courses/bisc441/Course_Materials/Readings/9-(Lect6)Crespi2000.pdf

345. Davis MB. 1990. Climate change and the survival of 

forest species. Chapter 6. In: Woodwell GM, editor. The Earth in 

Transition: Patterns and Processes of Biotic Impoverishment. 

Cambridge, United Kingdom: Cambridge University. p 99-110. 

Type: Book Section

Geographic Area: Global

Compilers’ Keywords: pollen, migration, paleoecology, trees, 

Tsuga canadensis

Introduction: The fossil record provides many examples of 

changes in geographical distributions of plants in response to 

changing climate. These examples provide a basis for predicting response to future climate. The accumulation of greenhouse 

gases in the atmosphere will soon cause climatic warming. 

With CO2 doubling, global temperatures are expected to rise 

3 degrees ± 1.5 °C degrees. If emission of greenhouse gases 

continues to increase at the present rates, global warming will 

average 0.3 °C per decade for the next century. Can the flora 

adjust easily? Or will climatic change cause extinction? The 

survival of many taxa through a series of glacial-interglacial 

cycles during the Pleistocene suggests that most plant species in the temperate zone have been able to disperse to keep 

pace with climatic changes in the past. Temperate zone trees 

survived the last glacial maximum as small populations in refugia in southeastern United States. As the ice sheets melted 

and the climate warmed at the end of the glacial interval, these 

plants were able to disperse northward hundreds of kilometers 

the aims, foci, and standards of natural resource management. 

Accordingly, the crucial project of natural resource law must 

be improving governance by cultivating agency accounting 

and learning to better manage uncertainty, promoting opportunities for interjurisdictional collaboration, and fostering public 

information and deliberation over the tradeoffs of strategies 

like assisted migration and the resource values that matter. 

343. Camacho A. 2013. The law and ethics of assisted migration. In: Assisted Migration: A Primer for Reforestation and 

Restoration Decision Makers; Portland, OR. 42 p.

Type: Presentation

Geographic Area: Global

Compilers’ Keywords: policies, EPA, Endangered Species 

Act, natural resource management

Abstract: To avoid extinctions and other harms to ecological health from escalating climatic change, scientists, resource 

managers, and activists are considering and even engaging 

in “assisted migration” – the intentional movement of an organism to an area in which its species has never existed. This 

article explores the profound implications of climate change 

for American natural resource management through the lens of 

this controversial adaptation strategy. It details arguments regarding the scientific viability and legality of assisted migration 

under the thicket of laws that govern natural resources in the 

United States. The article asserts, however, that the fundamental tensions raised by the strategy are ethical: to protect endangered species or conserve native biota; to manage ecological 

systems actively or leave nature wild and uncontrolled; and to 

preserve resources or manage them to promote their fitness under future conditions. The article explains why contemporary 

natural resource law’s fidelity to historic baselines, protecting 

preexisting biota, and shielding nature from human activity is 

increasingly untenable, particularly in light of climate change. 

Active, anticipatory strategies such as assisted migration may 

not only be permissible but even necessary to avert substantial irreversible harm to ecological systems. Scientists and 

resource managers should focus on developing scientific data 

to aid analyses of the risks and benefits of assisted migration 

in particular circumstances. To help develop such data while 

minimizing ecological harm, the article proposes provisionally 

limiting experimental translocations to situation where translocation is technically and economically feasible, and where the 

species is endangered, ecologically valuable, and compatible 

with the proposed site. 

Link: http://www.rngr.net/resources/assisted-migration

344. Crespi BJ. 2000. The evolution of maladaptation.

Heredity 84:623-629.

Type: Journal 

Geographic Area: Global
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 125

colonization will be inappropriate, especially when the broader 

social-ecological context of threatened species conservation is 

taken into account. 

Link: http://www.ncbi.nlm.nih.gov/pubmed/19577323

348. Hannah L. 2008. Protected areas and climate change. 

Annals of the New York Academy of Sciences 1134:201-212.

Type: Journal 

Geographic Area: Global

Keywords: protected areas, climate change, biodiversity, reserves, connectivity, corridors, assisted migration

Abstract: The study of protected areas and climate change 

has now spanned two decades. Pioneering work in the late 

1980s recognized the potential implications of shifting species range boundaries for static protected areas. Many early 

recommendations for protected area design were general, emphasizing larger protected areas, buffer zones, and connectivity 

between reserves. There were limited practical tests of these 

suggestions. Development of modeling and conservation planning methods in the 1990s allowed more rigorous testing of 

concepts of reserve and connectivity function in a changing 

climate. These studies have shown decreasing species representation in existing reserves due to climate change, and the 

ability of new protected areas to help slow loss of representation in mid-century scenarios. Connectivity on protected 

area periphery seems more effective than corridors linking 

protected areas. However, corridors serving other purposes, 

such as large carnivore movement, may be useful for accommodating species range shifts as well. Assisted migration and 

ex situ management strategies to complement protected areas 

are being explored. Finally, in scenarios of the latter half of the 

century, protected areas and connectivity become increasingly 

expensive and decreasingly effective, indicating the importance of reducing human-induced climate change.

349. Hewitt N, Klenk N, Smith AL, Bazely DR, 

Yan N, Wood S, MacLellan JI, Lipsig-Mumme C, 

Henriques I. 2011. Taking stock of the assisted migration 

debate. Biological Conservation 144:2560-2572.

Type: Journal 

Geographic Area: Global

Keywords: assisted migration, assisted colonization, managed relocation, climate change adaptation, debate

Abstract: Assisted migration was proposed several decades 

ago as a means of addressing the impacts of climate change 

on species populations. While its risks and benefits have been 

debated, and suggestions for planning and management given, 

there is little consensus within the academic literature over 

whether to adopt it as a policy. We evaluated the main features 

of the assisted migration literature including the study methods, taxonomic groups, geographic regions and disciplines 

to occupy their present-day range. Future greenhouse warming, however, differs from climatic changes in the past: the rate 

of change will be at least one order of magnitude more rapid. 

Examples from the most recent time of relatively rapid change 

exist—the warming just at the end of the last glacial interval 

in northwestern Europe fossils from warmth-requiring species 

of water plants and beetles are found earlier than the oldest 

fossils from trees. This suggests that the climate became warm 

enough for trees several centuries before trees began to grow 

on the local landscape. Dispersal of tree seeds was insufficient 

to track the change in temperature, or development of soils was 

not rapid enough to provide appropriate habitats for trees. 

346. Davis MB, Shaw RG. 2001. Range shifts and 

adaptive responses to Quaternary climate change. Science 

292:673-679.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: vegetation change, latitude shifts, 

elevation shifts, migration, extinction, tree

Abstract: Tree taxa shifted latitude or elevation range in response to changes in Quaternary climate. Because many modern trees display adaptive differentiation in relation to latitude 

or elevation, it is likely that ancient trees were also so differentiated, with environmental sensitivities of populations throughout the range evolving in conjunction with migrations. Rapid 

climate changes challenge this process by imposing stronger 

selection and by distancing populations from environments to 

which they are adapted. The unprecedented rates of climate 

changes anticipated to occur in the future, coupled with land 

use changes that impede gene flow, can be expected to disrupt 

the interplay of adaptation and migration, likely affecting productivity and threatening the persistence of many species.

Link: http://web.clas.ufl.edu/users/mbinford/GEOXXXX_

Biogeography/LiteratureForLinks/Davis_and_Shaw_2001_

range_shifts_climate_adaption_Science.pdf

347. Fazey I, Fischer J. 2009. Assisted colonization is a 

techno-fix. Trends in Ecology and Evolution 24:475.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: animals, conservation, assisted migration, ecosystem, biological extinction, biological models, 

risk assessment

Introduction: In this letter, authors agree with the tenet of 

the argument of Ricciardi and Simberloff (2009) that the attention and credibility given to such schemes is worrying and 

that it might send an overly optimistic message about assisted 

colonization to policymakers and the public. Here we propose 

three important additional considerations which lend further 

support to the view that widespread implementation of assisted 

CHAPTER 3 • MIGRATION • GENERAL
126 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

but I believe we should allocate a small portion of our attention 

to the issue of assisted colonization. McLachlan et al. propose 

framing the debate around two considerations—perception 

of risk and confidence in ecological understanding—that can 

be construed to generate an axis or continuum from scientists 

who would strongly support assisted colonization to those who 

would oppose it. I think it is useful to advance this exercise by 

considering three issues that can also be construed as continua: 

species that are more or less acceptable to translocate, sites that 

are more or less acceptable for receiving translocations, and 

projects that are more or less acceptable because of their socioeconomic ramifications and feasibility. I have used the term 

assisted colonization in contrast to assisted migration used by 

McLachlan et al. because many animal ecologists reserve the 

word migration for the seasonal, round-trip movements of animals and because the real goal of translocation goes beyond 

assisting dispersal to assuring successful colonization, a step 

that will often require extended husbandry.

Link: http://www.ncbi.nlm.nih.gov/pubmed/17883502

351. Iverson LR, Peters MP, Matthews S, Prasad 

AM. 2013. An overview of some concepts, potentials, issues, 

and realities of assisted migration for climate change adaptation 

in forests. In: Browning J, Palacios P, editors. Proceedings of 

the 60th Annual Western International Forest Disease Work 

Conference. Tahoe City (CA): USDA Forest Service, Northern 

Research Station. p 25-34.

Type: Government Document

Geographic Area: Global

Compilers’ Keywords: assisted migration, climate change, 

land management

Abstract: The climate has always been changing, but the 

rapid rate of climate change, as projected by the IPCC (2007) 

will likely place unique stresses on plant communities. In addition, anthropogenic barriers (e.g., fragmented land use) present a significant modern constraint that will limit the ability of 

species migration in responses to a changing climate. As such, 

managers are faced with four options that lay along a continuum when managing species in the face of climate change: (1) 

They can do nothing, and therefore allow existing landscapes 

to change without active intervention, accepting unknown or 

risky outcomes; (2) They can rely on passive resource management strategies to allow accommodation, such as Linking existing preserves with corridors; (3) They can actively manage 

landscapes to preserve them as they are, thus create refuges. 

Such habitat management would include actions like preventing invasions, installing irrigation, and regulating biotic interactions; or (4) They can actively manage landscapes to convert 

them into something deemed more compatible with projected 

climatic conditions. This last example of management would 

include assisted migration. The specific risks and benefits of 

each of these actions will depend upon the magnitude of climate 

involved. We further assessed the debate about the use of assisted migration, the main barriers to consensus, and the range of 

recommendations put forth in the literature for policy, planning 

or implementation. Commentaries and secondary literature reviews were as prevalent as first-hand scientific research and 

attention focused on a global rather than regional level. There 

was little evidence of knowledge transfer outside of the natural sciences, despite the obvious policy relevance. Scholarly 

debate on this topic has intensified during the last 3 years. We 

present a conceptual framework for evaluating arguments in 

the debate, distinguishing among the direct risks and benefits 

to species, ecosystems and society on the one hand, and other 

arguments regarding scientific justification, evidence-base and 

feasibility on the other. We also identify recommendations with 

potential to advance the debate, including careful evaluation of 

risks, benefits and trade-offs, involvement of relevant stakeholders and consideration of the complementarity among assisted migration and less risk tolerant strategies. We conclude, 

however, that none of these will solve the fundamental, often 

values-based, challenges in the debate. Solutions are likely to 

be complex, context-dependent and multi-faceted, emerging 

from further research, discussion and experience.

Link: http://www.researchgate.net/publication/232361203_

Taking_stock_of_the_assisted_migration_debate/

file/79e41512f9fdb8e5a8.pdf

350. Hunter ML, Jr. 2007. Climate change and moving species: furthering the debate on assisted colonization.

Conservation Biology 21:1356-1358.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: animals, conservation, biological 

extinction, greenhouse effect, population dynamics, assisted 

migration

Introduction: With global climate change looming large in 

the public psyche, the recent paper by McLachlan et al. (2007) 

and its popular accompaniment (Fox 2007) are timely indeed. 

Of course some conservation biologists will not wish to think 

about the prospect of actively moving species that are threatened with extinction by climate change. For them this would 

be almost analogous to handing out placebos in the midst of 

an epidemic and worse yet, these placebos may have serious 

unintended consequences if translocated species become invasive. They will probably argue that we should focus almost 

exclusively on two central roles for conservation biology: (1) 

facilitating natural range shifts by redoubling efforts to maintain or restore large scale connectivity and (2) working with 

our fellow environmental professionals to avoid carbon-management solutions that will have unacceptable consequences 

for biodiversity (e.g., by directing biofuel production away 

from sites that would involve the conversion of native vegetation into fuel farms. These two roles will be very demanding, 

CHAPTER 3 • MIGRATION • GENERAL
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 127

CHAPTER 3 • MIGRATION • GENERAL

(common garden /provenance trials) to assist with making seed 

movement guidelines and research is needed.

Link: http://nativeseed.info/2010/session/climate.html#kaye

353. Keim B. 2008. Last-ditch resort: move polar bears to 

Antarctica? Wired 17 July 2008. (URL accessed 25 November 

2015)

Type: Article

Geographic Area: Global

Compilers’ Keywords: migration, conservation, extinctions

Introduction: It may seem like a preposterous question. But 

polar bears are just the tip of the “assisted colonization” iceberg. 

Other possibilities: moving African big game to the American 

Great Plains, or airlifting endangered species from one mountaintop to another as climate zones shrink. Once dismissed as 

wrongheaded and dangerous, assisted colonization—rescuing 

vanishing species by moving them someplace new—is now 

being discussed by serious conservationists. And no wonder: 

Caught between climate change and human pressure, species 

are going extinct 100 times faster than at any point in human 

history. And some scientists say that figure is too conservative. 

The real extinction rate, they say, is a full 1,000 times higher 

than normal. The last time such annihilation took place was 

during the time of the dinosaurs. And though many conservationists say that saving species by transplanting them is foolish, 

others say there’s no choice.

Link: http://www.wired.com/science/planetearth/

news/2008/07/species_relocation?currentPage=all

354. Koskela J, Vinceti B, Dvorak W, Bush D, 

Dawson IK, Loo J, Kjaer ED, Navarro C, Padolina C, 

Bordács S, Jamnadass R, Graudal L, Ramamonjisoa 

L. 2014. Utilization and transfer of forest genetic resources: a 

global review. Forest Ecology and Management 333:22-34.

Type: Journal 

Geographic Area: Global

Keywords: reproductive material, plantations, introduced 

species, access and benefit sharing, Nagoya Protocol

Abstract: Over the last 200 years, genetic resources of forest 

trees have been increasingly transferred, within and outside of 

species’ native distribution ranges, for forestry and for research 

and development (R&D). Transferred germplasm has been deployed to grow trees for numerous purposes, ranging from the 

production of wood and non-wood products to the provision 

of ecosystem services such as the restoration of forests for biodiversity conservation. The oldest form of R&D, provenance 

trials, revealed early on that seed origin has a major influence 

on the performance of planted trees. International provenance 

trials have been essential for selecting seed sources for reforestation and for improving tree germplasm through breeding. 

Many tree breeding programmes were initiated in the 1950s, 

pressure, the context of the ecosystem and its landscape, and 

the goals of human decisions. This paper describes some options on how to decide among the above choices, introduces 

assisted migration, and describes the possible ramifications 

associated with it. We then present one research approach to 

assist in locating and evaluating potential applications of assisted migration.

Link: http://www.treesearch.fs.fed.us/pubs/45280

352. Johnson R. 2010. Seed movement and climate change 

from a Forest Service perspective. In: National Native Seed 

Conference: Native Plant Materials Development, Production, 

and Use for Habitat Restoration. Corvallis (OR): Institute for 

Applied Ecology. 39 p.

Type: Presentation

Geographic Area: USA

Compilers’ Keywords: forest genetics, forest management, 

seedlot selection tool

Abstract: The USDA Forest Service (USFS) Native plant restoration policy calls for use of “genetically appropriate” native 

plant material on USFS lands. This is typically accomplished 

by delineating seed zones to restrict seed movement to ensure 

locally-adapted plant material is used in restoration / reforestation / rehabilitation efforts. Given that the climate is changing 

more rapidly than any time in the recent past, our starting point 

of “local is best” may not be the optimal option given that local climates have changed over the last few decades and even 

more change is forecasted in the future. While the USFS policy has not changed, USFS Regional Geneticists, along with 

USFS researchers, came together for a 3-day workshop to discuss seed movement options in light of climate change (a paper 

is forthcoming). The workshop centered on forest trees, not 

smaller native plants. Discussions acknowledged that climate 

has already changed in the last few decades (surface temperature has warmed about 1 °F) and that certain forest tree species 

have migrated in some places. Both genetic theory and species migration models suggest that climate change will impact 

species differently. The impact of climate change on a species 

will be function of lifespan, mating system, dispersal capabilities, habitat specialization, genetic variation, population size, 

pest/pathogen interactions, etc. Key points coming out of the 

meeting included: (1) The importance of deploying genetically 

diverse populations in our restoration efforts. (2) Local sources 

still appear to be appropriate choices at present, but including additional sources from adjacent seed zones could provide 

the variation needed for the future. (3) At least one region is 

moving seed up one elevation or “north” one cold hardiness 

band. (4) We will take minimal risks over large scales and 

larger risks at smaller scales (i.e., only do assisted migration 

studies over large distances on experimental scales). (5) The 

need for gene conservation programs is increasing in light of 

climate change. (6) For most species, we have no genetic data 
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CHAPTER 3 • MIGRATION • GENERAL

frameworks and numerous recommendations for weighing and 

evaluating these factors. Here, we argue that this debate is, in 

part, misguided, and that a more productive discussion would 

result if the issue were reframed in light of (1) the goals of 

assisted colonization, (2) the realities of projected climate impacts, and (3) the use of complementary adaptation strategies, 

such as enhancing landscape connectivity.

Link: http://depts.washington.edu/oldenlab/wordpress/wpcontent/uploads/2013/03/FrontiersEcoEnv_2011.pdf

356. McLachlan JS, Hellmann JJ, Schwartz MW. 

2007. A framework for debate of assisted migration in an era of 

climate change. Conservation Biology 21:297-302.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: conservation, demography, ecosystem, biological extinction, genetic variation, greenhouse effect, 

biological models, population dynamics, policy, framework

Introduction: Assisted migration is contentious issue that 

places different conservation objectives at odds with one another. This element of debate, together with the growing risk 

of biodiversity loss under climate change, means the now is 

the time for the conservation community to consider assisted 

migration. Our intent here is to highlight the problem caused 

by a lack of a scientifically based policy on assisted migration, 

suggest a spectrum of policy options, and outline a framework 

for moving toward a consensus on this emerging conservation 

dilemma. 

Link: http://www.ncbi.nlm.nih.gov/pubmed/17391179

357. Millar CI. 2013. The role of assisted migration in 

climate adaptation planning: when and where to employ it? In: 

Assisted Migration: A Primer for Reforestation and Restoration 

Decision Makers; Portland, OR. 42 p.

Type: Presentation

Geographic Area: Global

Compilers’ Keywords: assisted migration, climate change, 

natural migration, decision making, resistance, resilience, 

response, realignment, spatial scales, temporal scales, 

paleoclimate

Abstract: In the face of changing climates, either historic or 

contemporary, species respond by adapting, moving, or dying 

(going extinct or populations extirpating). Assisted migration 

commonly has been discussed in the context of genetic adaptation. As background to addressing the questions of “When 

and where to employ assisted migration?” I briefly review 

as well the ways in which species have moved in response 

to historic climate change. This begins with an overview of 

principles regarding natural climate change that are relevant 

to decision-making for contemporary climate adaptation. In an 

historic context, climates: 1) have been changing continually; 

but as one round of testing and selection typically takes decades, the most advanced of them are only in their third cycle. 

Recent advances in forest genomics have increased the understanding of the genetic basis of different traits, but it is unlikely 

that molecular marker-assisted approaches will quickly replace 

traditional tree breeding methods. Furthermore, provenance 

trials and progeny tests are still needed to complement new 

research approaches. Currently, seed of boreal and temperate 

trees for reforestation purposes are largely obtained from improved sources. The situation is similar for fast growing tropical and subtropical trees grown in plantations, but in the case of 

tropical hardwoods and many agroforestry trees, only limited 

tested or improved seed sources are available. Transfers of tree 

germplasm involve some risks of spreading pests and diseases, 

of introducing invasive tree species and of polluting the genetic make-up of already present tree populations. Many of these 

risks have been underestimated in the past, but they are now 

better understood and managed. Relatively few tree species 

used for forestry have become invasive, and the risk of spreading pests and diseases while transferring seed is considerably 

lower than when moving live plants. The implementation of 

the Nagoya Protocol on access to genetic resources and benefit sharing (ABS) may significantly change current transfer 

practices in the forestry sector by increasing transaction costs 

and the time needed to lawfully obtain forest genetic resources 

for R&D purposes. Many countries are likely to struggle to 

establish a well-functioning ABS regulatory system, slowing 

down the process of obtaining the necessary documentation for 

exchange. This is unfortunate, as climate change, outbreaks of 

pests and diseases, and continual pressure to support productivity, increase the need for transferring tree germplasm and 

accelerating R&D.

Link: http://www.bioversityinternational.org/uploads/

tx_news/Utilization_and_transfer_of_forest_genetic_

resources_a_global_review_1869.pdf

355. Lawler JJ, Olden JD. 2011. Reframing the debate over assisted colonization. Frontiers in Ecology and the 

Environment 9:569-574.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: migration, climate change, landscape 

connectivity, managed relocation, framework, guidelines

Abstract: Assisted colonization—also known as managed relocation or assisted migration—is one way of facilitating range 

shifts for species that are restricted in their ability to move in 

response to climate or other environmental changes. Over 

the past decade, a healthy debate has evolved in the scientific 

community over the costs and benefits of assisted colonization as a climate-adaptation strategy. This discussion has focused largely on the specific risks and benefits of intentionally 

moving species, and has led to the development of multiple 
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CHAPTER 3 • MIGRATION • GENERAL

regulatory opportunities and constraints relative to implementing assisted migration. 

Link: http://www.rngr.net/resources/assisted-migration

358. Minteer BA, Collins JP. 2010. Move it or lose it? 

The ecological ethics of relocating species under climate change. 

Ecological Applications 20:1801-1804.

Type: Journal 

Geographic Area: Global

Keywords: climate change, ecological policy, ethics, managed relocation, species conservation

Abstract: Managed relocation (also known as assisted colonization, assisted migration) is one of the more controversial 

proposals to emerge in the ecological community in recent 

years. A conservation strategy involving the translocation of 

species to novel ecosystems in anticipation of range shifts 

forced by climate change, managed relocation (MR) has divided many ecologists and conservationists, mostly because 

of concerns about the potential invasion risk of the relocated 

species in their new environments. While this is indeed an important consideration in any evaluation of MR, moving species across the landscape in response to predicted climate shifts 

also raises a number of larger and important ethical and policy 

challenges that need to be addressed. These include evaluating the implications of a more aggressive approach to species 

conservation, assessing MR as a broader ecological policy 

and philosophy that departs from longstanding scientific and 

management goals focused on preserving ecological integrity, 

and considering MR within a more comprehensive ethical and 

policy response to climate change. Given the complexity and 

novelty of many of the issues at stake in the MR debate, a more 

dynamic and pragmatic approach to ethical analysis and debate 

is needed to help ecologists, conservationists, and environmental decision makers come to grips with MR and the emerging 

ethical challenges of ecological policy and management under 

global environmental change.

Link: http://cirge.stanford.edu/Minter_managedrelocation_

ethics_2010.pdf

359. Odenbaugh J. 2013. Assisted migration and invasive 

species: exploring an ethical dilemma. In: Assisted Migration: 

A Primer for Reforestation and Restoration Decision Makers; 

Portland, OR. 22 p.

Type: Presentation

Geographic Area: Global

Compilers’ Keywords: ecological morality, species extinctions, climate change, exotic species

Abstract: According to many, we are subject to two duties. 

First, it is morally wrong for human to cause a species to go 

extinct. Second, it is morally wrong to introduce a species into 

an area in which it is not native. Unfortunately, human-induced 

2) express at multiple and nested scales, including interannual, 

decadal, centennial, and millennial; 3) are driven by mostly 

independent mechanisms at each scale; and 4) cumulatively 

result in climate changes that have been novel, gradual and 

directional, episodic and quasi-reversible, and chaotic. At long 

time scales and in response to large climate changes, species 

have migrated (shifted ranges) long distances (e.g., hundreds 

of kilometers) in regions of low relief or across large elevation 

gradients (e.g., 1000 m) in mountainous environments. Species 

responded individualistically based on life-history characteristics and inherent climate envelopes. As a result, community 

compositions (species diversity) were transient, emerging as 

novel assemblages of species associated together at any one 

time and place. In response to shorter time-scale and smaller 

magnitude climate changes, species responded less often with 

biogeographic range contractions or expansions and more 

through changes in demography, species dominance relationships, density and vigor, and disturbance regimes. Lessons 

that emerge for climate adaptation focus on working with, not 

against, these inherent adaptive responses of species in the face 

of climate change. That change has been constant, hoever, does 

not imply that “anything goes.” I review lessons learned from 

natural climate responses for implications to concepts of native 

range, neo-native distributions, community resilience, novel 

climates and non-analog associations, “red queen” responses, 

the role of climatic refugia in maintenance of diversity, and 

pace of change following disturbance. With this background, I 

turn to issues regarding decision—making for assisted migration: when and where to employ it in the face of contemporary 

climate change. Institutional context influences these factors, 

whether goals are primarily for ecosystem services (production of goods and services) or ecosystem management (species protection and recovery). First principles apply, including: do no harm, don’t fix what isn’t broken, learn as you go; 

leave intact escape routes; and weigh risk and urgency into 

decisions. Scale also affects decisions regarding “When?” in 

climate adaptation: actions justifiably made for the short term 

or small spatial scale are likely different from those focused 

on long-term investments and benefits. In regard to “Where?” 

considerations are important for both the source (donor) and 

destination locations. In the former, genetic concerns often are 

paramount, but ecological issues such as co-factors of disease 

or insect transmission, history of the population, and vigor of 

propagules (seed or ramet stock) are equally important. Factors 

to consider at the destination locations include concerns for 

both the target species (how well it will thrive) and the receiving ecosystem (how it is affected). Effects to weight include 

invasiveness, genetic contamination, allelo-chemical and soilnutrient alterations, phenology, community interactions and 

species displacement, and influences on disturbance regimes. 

I review these considerations relative to the four fundamental 

options for climate adaptation planning: resistance, resilience, 

response, and realignment (restoration), and provide illustrative examples for each. In conclusion I comment on policy and 
130 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

CHAPTER 3 • MIGRATION • GENERAL

conceptions of AM have not been clearly distinguished in the 

literature, which has added confusion to recent dialogue and 

debate. Here, we aim to facilitate clearer communication on 

this ftopic by detailing this distinction and encouraging a more 

nuanced view of AM.

362. Peters RL. 1992. Conservation of biological 

diversity in the face of climate change. Chapter 2. In: Peters RL, 

Lovejoy TE, editors. Global warming and biological diversity. 

New Haven (CT): Yale University Press. p 15-31. 

Type: Book Section

Geographic Area: Global

Compilers’ Keywords: climate change, range shifts, conservation, guidelines

Introduction: We can infer how the biota might respond 

to climate change by observing present and past distributions of plants and animals, which are largely determined by 

temperature and moisture patterns. For example, one race of 

dwarf birch (Betula nana) can grow only where the temperature never exceeds 22 °C, suggesting that it would disappear 

from those areas where global warming causes temperatures 

to exceed 22 °C. Recent historical observations of changes in 

range and species dominance, such as the gradual replacement 

spruce (Picea rubens) by deciduous species during the past 

180 years in the eastern United States, can also suggest future 

responses. Insight into long-term responses to large climatic 

changes can be gleaned from studies of fossil distributions of, 

particularly, pollen and small mammals. Such observations tell 

us that plants and animals are very sensitive to climate. Their 

ranges move when the climate patterns change—species die 

out in areas where they were once found and colonize new 

areas where the climate becomes newly suitable. 

363. Peters RL, Darling JDS. 1985. The greenhouseeffect and nature reserves. BioScience 35:707-717.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: assisted migration, climate change, 

biodiversity, global warming, conservation, review, species 

extinctions

Introduction: Current human development and population 

trends suggest to all but the very optimistic that by the next 

century most other surviving terrestrial species may well be 

relegated to small patches of their original habitat, patches isolated by vast areas of human-dominated urban and agricultural 

lands. Without heroic measures of habitat conservation and 

intelligent management, hundreds of thousands of plant and 

animal species could become extinct by the end of this century, 

with more to follow in the next. In this paper, we will identify 

problems caused by climate change that affect biological communities, examine the particular difficulties faced by species 

climate change will cause species to go extinct unless we relocate those species to areas outside their native range. Thus, we 

are either causing species to go extinct or creating exotic species both of which are morally wrong. In this talk, I consider 

ways of dealing with the environmental dilemma. 

Link: http://www.rngr.net/resources/assisted-migration

360. Park A, Talbot C. 2012. Assisted migration: 

uncertainty, risk and opportunity. The Forestry Chronicle 

88:412-419.

Type: Journal 

Geographic Area: Canada

Keywords: assisted migration, managed relocation, climate 

change scenarios, uncertainty, ecophysiology, epigenetics

Abstract: The recent Forestry Chronicle special section on 

the subject of Assisted Migration (AM) did a great service to 

the Canadian forestry community by summarizing the risks, 

opportunities and ecological aspects of this forest management 

strategy. In this paper, we expand on some aspects of AM that 

were discussed in the special section, as well as discussing 

additional dimensions of AM that should be considered and 

debated. We expand on the theme of scientific uncertainties 

around future warming, emphasizing the full extent of uncertainty in estimates of climate sensitivity. We also expand upon 

and summarize a number of recent ecophysiological results 

that have implications for the adaptation and acclimation of 

trees to climate change. We also discuss opportunities for innovative forest management, the influence of economic trends 

on the future of the Canadian forest industry, and limitations on 

public knowledge of climate change, all of which are factors 

that will influence the feasibility of AM schemes in the future.

361. Pedlar J, McKenney DW, Aubin I, Beardmore 

T, Beaulieu J, Iverson LR, O’Neill GA, Winder RS, 

Ste-Marie C. 2012. Placing forestry in the assisted migration debate. BioScience 62:835-842.

Type: Journal 

Geographic Area: North America

Keywords: assisted migration, climate change, forestry, conservation, trees 

Abstract: Assisted migration (AM) is often presented as a 

strategy to save species that are imminently threatened by rapid 

climate change. This conception of AM, which has generated 

considerable controversy, typically proposes the movement 

of narrowly distributed, threatened species to suitable sites 

beyond their current range limits. However, existing North 

American forestry operations present an opportunity to practice AM on a larger scale, across millions of hectares, with a 

focus on moving populations of widely distributed, nonthreatened tree species within their current range limits. Despite 

these differences (and many others detailed herein), these two 
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CHAPTER 3 • MIGRATION • GENERAL

environments. Few long-term field studies have been done. 

The critical question to Davis and a colleague, Catherine 

Zabinski, is how quickly trees can migrate, for the fate of numerous species in North America will depend on whether they 

can shift north to cooler climates when their current range becomes uninhabitable. 

366. Safford HD, Hellmann JJ, McLachlan JM, Sax 

DF, Schwartz MW. 2009. Managed relocation of species: 

Noah’s Ark or Pandora’s Box? Eos 90:1.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: assisted migration, species 

conservation

Introduction: “Managed relocation” (MR; also called “assisted migration” or “assisted colonization”) is the human-aided movement of species adversity affected by global change 

where these species cannot move themselves. Goals of MR 

include, but are not limited to, conservation of biodiversity, reduction of extinction risk, enhancement of evolutionary potential, and maintenance or augmentation of ecosystem services. 

Recently there has been much talk, in the scientific literature 

as well as in the press, about the costs and benefits of MR. 

As a conservation strategy, MR has promise, but it could also 

have serious collateral costs. For example, MR may succeed 

in rescuing a given species from extinction, but it may also 

introduce a species into habitat where it becomes invasive, 

causing ecosystem disruptions or extinctions of other taxa. A 

working group met in August 2008, in conjunction with the 

annual meeting of the ESA, to develop a framework for understanding the degree to which MR could achieve its objectives, 

the risks that it might incur, and strategies that could be used 

to implement it. 

367. Sandler R. 2010. The value of species and the ethical foundations of assisted colonization. Conservation Biology 

24:424-431.

Type: Journal 

Geographic Area: Global

Keywords: assisted colonization, biodiversity, ecosystem 

management, value of species

Abstract: Discourse around assisted colonization focuses on 

the ecological risks, costs, and uncertainties associated with the 

practice, as well as on its technical feasibility and alternative 

approaches to it. Nevertheless, the ethical underpinnings of 

the case for assisted colonization are claims about the value of 

species. A complete discussion of assisted colonization needs 

to include assessment of these claims. For each type of value 

that species are thought to possess it is necessary to determine 

whether it is plausible that species possess the type of value 

and, if so, to what extent their possessing it justifies assisted 

in biological reserves, and suggest management options. 

Although we recognize that dealing with short-term extinction threats alone will strain the resources of conservationists, 

we feel that the possible negative effects of global warming 

could be so severe that conservation plans should be amended 

to reflect knowledge of climatic effects as soon as it becomes 

available. Decisions about the siting and design of reserves and 

assumptions about how much management will be needed in 

the future must reflect the increased demands, both economic 

and biological, of global warming. 

364. Ricciardi A, Simberloff D. 2009. Assisted colonization is not a viable conservation strategy. Trends in Ecology 

and Evolution 24:248-253.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: animals, conservation, ecosystem, 

biological extinction, human activities, biological models, 

plants, assisted migration, management

Abstract: A potential conservation strategy increasingly discussed by conservation biologists is the translocation of species 

to favorable habitat beyond their native range to protect them 

from human-induced threats, such as climate change. Even if 

preceded by careful risk assessment, such action is likely to 

produce myriad unintended and unpredictable consequences. 

Accurate risk assessment is impeded by contingency: the impacts of introduced species vary over time and space under the 

influence of local environmental variables, interspecific interactions and evolutionary change. Some impacts, such as native 

species extinctions, are large and irrevocable. Here we argue 

that conservation biologists have not yet developed a sufficient 

understanding of the impacts of introduced species to make 

informed decisions regarding species translocations.

Link: http://www.ncbi.nlm.nih.gov/pubmed/19324453

365. Roberts L. 1989. How fast can trees migrate?

Science 243:735-737.

Type: Journal 

Geographic Area: USA

Compilers’ Keywords: research news, tree migration, climate change, forest management, review, rates

Introduction: The forests in the south will go first. Seedlings 

will wither, the understory plants will be replaced. And over 

the next century or so, many now-abundant trees will go extinct across much of the United States. Such are the predictions of Margaret Davis, one of handful of ecologists looking 

at the effects of the much discussed greenhouse warming on 

North American forests. As Davis and others who make these 

predictions readily admit, they are riddled with uncertainties. 

Data are scarce and have been cobbled together from fossil 

records, theoretical models, and experiments in controlled 
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CHAPTER 3 • MIGRATION • GENERAL

369. Schlaepfer MA, Helenbrook WD, Searing 

KB, Shoemaker KT. 2009. Assisted colonization: evaluating contrasting management actions (and values) in the face of 

uncertainty. Trends in Ecology and Evolution 24:421-422.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: assisted migration, conservation, 

debate

Introduction: In their recent opinion article in Trends in 

Ecology & Evolution, Ricciardi and Simberloff argue that assisted colonization is not an appropriate management option 

because the impacts of introduced species are too difficult to 

predict, and can have harmful consequences for recipient ecosystems, including the extinction of native species. Invoking 

the precautionary principle, the authors argue that alternative 

conservation tactics must be pursued, even for species faced 

with extinction in their native range. We concur with the authors that there are risks associated with introducing species 

outside of their current range. However, we disagree that these 

risks are so great that assisted colonization should not be considered among possible management options.

Link: http://noss.cos.ucf.edu/papers/Schlaepfer%20et%20

al%202009%20and%20Sax%20et%20al%202009.pdf

370. Schwartz MW. 1994. Conflicting goals for conserving biodiversity: issues of scale and value. Natural Areas Journal 

14:213-216.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: review, conservation, translocation, 

assisted migration

Abstract: Differing conservation values create a multiplicity of goals toward which conservation activities are directed. 

Multiple and differing goals also are inherent in conservation 

at differing spatial and temporal scales. In many cases these 

different goals result in management actions that are complementary and that simultaneously benefit many species and 

habitats. In contrast, there are instances in which conservation 

for one set of values, or at one spatial scale, suggests a management action that would violate other conservation goals. I 

define the goals and value systems used to support biological 

conservation and present three cases where differing conservation objectives conflict. Specifically, I address how (1) a 

proposed translocation of an endangered species outside its 

historic range, to prevent its extinction, would violate the historic integrity of the recipient community; (2) the use of fire, 

to maintain plant community composition in grasslands, may 

threaten native insect biodiversity; and (3) assisting the process of long-distance seed dispersal, to aid plant-range shifts 

disrupted by anthropogenic climate warming, would violate 

the integrity of recipient plant communities. While examples 

colonization. I conducted such an assessment for each of the 

predominant types of value ascribed to species: ecological, 

instrumental (including option value), existence, and intrinsic 

value (including interest-based, objective, and value-dependent 

intrinsic value). The vast majority of species, including several 

that have been proposed as candidates for assisted colonization, 

have much less value than is often presumed. Moreover, with 

respect to some types of value, assisted colonization would not 

fully preserve the value of the target species even if it were to 

keep the target species in existence. Therefore, the case for assisted colonization is significantly weaker and more qualified 

than its advocates often suppose. There may be exceptional 

species for which assisted colonization is well justified—and 

for this reason, case-by-case assessment is necessary—but in 

general the burden of justification generated by the ecological 

risks associated with assisted colonization is not met by the 

value potentially preserved by assisted colonization. This suggests that assisted colonization ought to have, at most, a very 

minor role in the portfolio of ecosystem management practices, even as it pertains to species conservation under conditions 

of rapid climate change.

Link: http://www.ncbi.nlm.nih.gov/pubmed/19878237

368. Sax DF, Smith KF, Thompson AR. 2009. 

Managed relocation: a nuanced evaluation is needed. Trends in 

Ecology and Evolution 24:472.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: assisted migration, climate change

Introduction: Managed relocation (aka ‘assisted colonization’ and ‘assisted migration’) aims to save species from the 

effects of climate change by purposefully transporting them to 

areas where they have not previously occurred, but where they 

are expected to survive as temperatures increase. In a recent 

Opinion article in TREE, Ricciardi and Simberloff suggest that 

‘assisted colonization is tantamount to ecological roulette and 

should probably be rejected as a sound conservation strategy 

by the precautionary principle.’ We disagree for three primary 

reasons. First, the precautionary principle is not a stand-alone 

reason to rule out managed relocation. Second, we know 

more about the impacts of species invasions than Ricciardi 

and Simberloff suggest, particularly with respect to species 

extinction. Third, because extinctions are permanent and irreversible, using managed relocation to reduce extinctions at 

the cost of changing the composition and functioning of ecosystems is a tradeoff that some managers might be willing to 

make. Ultimately, the risk of species extinctions from climate 

changes is too large to summarily discount managed relocation 

without first carefully evaluating its benefits and dangers in a 

nuanced way. 

Link: http://www.brown.edu/Research/Sax_Research_Lab/

Documents/PDFs/reply%20to%20managed%20relocation.pdf
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 133

used as a conservation tool to overcome barriers to dispersal. 

A dichotomy exists for conservation-oriented translocations: 

on one hand, there are those that release plants or animals into 

known historic ranges and on the other hand, there are releases 

outside historic distributions. Misuse of or attempts to redefine 

established terms and a proliferation of variants of new terms 

such as assisted colonization confuse and hamper communication. The aim of this opinion article is to describe and define 

a conservation translocation spectrum, from species reintroductions to assisted colonization, and beyond, and in so doing 

provide a standard framework and terminology for discussing 

translocation options. I suggest that we are moving along this 

spectrum, away from the dictates of historical species distribution records, toward the inclusion of more risky interventions that will be required to respond to habitat shifts due to 

anthropogenic impacts. To some extent rapid climate change 

changes everything, including how we should view introductions versus reintroductions. We need to seriously consider 

adding other approaches to our conservation toolbox. Assisted 

colonization will start us along this path, acknowledging as it 

does the accelerated rate of habitat change and the problems of 

attempting to preserve dynamic systems. The next step along 

the conservation translocation spectrum may be for reintroduction biology and restoration ecology to more comprehensively 

join forces on carefully selected projects to use species introductions to create novel ecosystems through active ecological 

community construction.

373. Shirey PD, Lamberti GA. 2010. Assisted colonization under the U.S. Endangered Species Act. Conservation 

Letters 3:45-52.

Type: Journal 

Geographic Area: USA

Keywords: assisted colonization, assisted migration, climate 

change, endangered species act, endangered species policy, 

experimental population provision, managed relocation, 

Mitchell’s satyr butterfly, Neonympha mitchellii mitchellii, 

translocation

Abstract: Assisted colonization could help prevent the extinction of threatened and endangered species by intentionally 

moving a species to a region where it has not occurred in the 

recent past, but should survive under future climate scenarios. 

Where species are naturally localized and confined to patchy 

habitats, assisted colonization might be the only means for 

population dispersal across human landscape barriers such as 

urban and agriculture areas. The major risk associated with assisted colonization is introducing ecologically harmful species. 

Previous policy papers have described management options for 

deciding when to move a species to mitigate for climate change. 

We build on this previous work by examining management options and policy solutions for assisted colonization under the 

U.S. Endangered Species Act (ESA). On its surface, the ESA 

statutory language appears to provide the legal framework for 

of conflicting goals are not hard to describe, they are hard to 

resolve and pose challenges to conservation biology that are 

not adequately addressed at the present time. Increased use of 

goal-setting would improve our ability to explicitly measure 

the success of conservation projects. Failure to set priorities 

and goals in conservation management implies a misplaced 

faith in the balance of nature. 

371. Schwartz MW, Hellmann JJ, McLachlan 

JM, Sax DF, Borevitz JO, Brennan J, Camacho 

AE, Ceballos G, Clark JR, Doremus H, Early R, 

Etterson JR, Fielder D, Gill JL, Gonzalez P, Green N, 

Hannah L, Jamieson DW, Javeline D, Minteer BA, 

Odenbaugh J, Polasky S, Richardson DM, Root TL, 

Safford HD, Sala O, Schneider SH, Thompson AR, 

Williams JW, Vellend M, Vitt P, Zellmer S. 2012.

Managed relocation: integrating the scientific, regulatory, and 

ethical challenges. BioScience 62:732-743.

Type: Journal 

Geographic Area: Global

Keywords: ethics, policy, law, conservation, translocation

Abstract: Managed relocation is defined as the movement of 

species, populations, or genotypes to places outside the areas 

of their historical distributions to maintain biological diversity 

or ecosystem functioning with changing climate. It has been 

claimed that a major extinction event is under way and that 

climate change is increasing its severity. Projections indicating that climate change may drive substantial losses of biodiversity have compelled some scientists to suggest that traditional management strategies are insufficient. The managed 

relocation of species is a controversial management response 

to climate change. The published literature has emphasized 

biological concerns over difficult ethical, legal, and policy issues. Furthermore, ongoing managed relocation actions lack 

scientific and societal engagement. Our interdisciplinary team 

considered ethics, law, policy, ecology, and natural resources 

management in order to identify the key issues of managed 

relocation relevant for developing sound policies that support 

decisions for resource management. We recommend that government agencies develop and adopt best practices for managed relocation.

372. Seddon PJ. 2010. From reintroduction to assisted 

colonization: moving along the conservation translocation spectrum. Restoration Ecology 18:796-802.

Type: Journal 

Geographic Area: Global

Keywords: ecological engineering, novel ecosystems, population restoration, species introductions

Abstract: Translocation, the intentional movement of living organisms from one area to another is increasingly being 

CHAPTER 3 • MIGRATION • GENERAL
134 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

in the scientific community and society at large, is growing 

rapidly and is starting to be translated into action in Canada. 

However, the concept is in its infancy; clear terminology has 

not yet been established and assisted migration still encompasses a broad range of practices. This introductory paper for 

the special issue of The Forestry Chronicle on the subject of assisted migration describes increasing interest in the subject and 

its complexity. It also provides an overview of the potential 

scale of assisted migration, proposes a terminology, and briefly 

introduces the following papers. Overall, the five papers aim to 

present a comprehensive state of the scientific and operational 

knowledge and the debate on assisted migration in the context 

of Canada’s forests.

376. Stone R. 2010. Home, home outside the range?

Science 329:1592-1594.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: assisted migration, climate change, 

conservation, assisted colonization, butterfly, Orchidaceae

Compilers’ Summary: One of the earlier popular press 

articles about moving species to new places in response to 

climate change. In China, orchids will be impacted by an increase in temperature, such that some scientists are considering 

assisted colonization to save species. But other scientists fear 

that movements may cause more harm than good. The article 

discusses both sides of the debate.

Link: http://plantecology.syr.edu/conbio/readings/Stone2010.

pdf

377. Thomas CD. 2011. Translocation of species, climate 

change, and the end of trying to recreate past ecological communities. Trends in Ecology Evolution 26:216-221.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: biota, conservation, environment, 

conservation, environment, extinction, review

Abstract: Many of the species at greatest risk of extinction 

from anthropogenic climate change are narrow endemics that 

face insurmountable dispersal barriers. In this review, I argue 

that the only viable option to maintain populations of these 

species in the wild is to translocate them to other locations 

where the climate is suitable. Risks of extinction to native species in destination areas are small, provided that translocations 

take place within the same broad geographic region and that 

the destinations lack local endemics. Biological communities 

in these areas are in the process of receiving many hundreds of 

other immigrant species as a result of climate change; ensuring 

that some of the ‘new’ inhabitants are climate-endangered species could reduce the net rate of extinction.

Link: http://www.ncbi.nlm.nih.gov/pubmed/21411178

CHAPTER 3 • MIGRATION • GENERAL

allowing assisted colonization, as the U.S. Congress gave the 

U.S. Fish and Wildlife Service (USFWS) broad discretion to 

manage populations of endangered species. However, current 

USFWS regulations are an impediment to assisted colonization for many endangered animal species, whereas regulations 

do not necessarily restrict assisted colonization of endangered 

plants. Because this discrepancy, we recommend a review of 

the regulatory language governing movements of endangered 

species.

Link: http://dev.cakex.org/sites/default/files/Assisted%20

colonization%20under%20the%20U.S.%20Endangered%20

Species%20Act.pdf

374. Sollenberger D, Vitt P, Yates E. 2010. Climate 

change, seeding banking and assisted migration - the long shadows of humanity. In: National Native Seed Conference: Native 

Plant Materials Development, Production, and Use in Habitat 

Restoration. Corvallis (OR): Institute for Applied Ecology. 33 

p.

Type: Presentation

Geographic Area: USA

Compilers’ Keywords: MaxEnt, Cirsium pitcheri, 

Andropogon gerardii, seed collection

Abstract: Rapid climate change has the potential to alter the 

climatic regime of many plant species globally. Plant species 

will respond variously via phenotypic plasticity, evolutionary 

adaptation, migration, or extinction. Using Species Distribution 

Model to predict how the bioclimatic envelopes for individual 

species might shift can provide insight into conservation strategies, including seed banking. When fragmentation limits the 

migration potential or when natural migration rates are outstripped by the pace of climate change, some propose purposeful, human-mediated migration (assisted migration or manage 

relocation) as a solution. We present a strategy to collect and 

bank seeds of plant species at risk of extinction in the face of 

rapid climate change to ensure that emerging habitats are as 

species-diverse as possible.

Link: http://nativeseed.info/2010/presentations/Vitt.pdf

375. Ste-Marie C, Nelson EA, Dabros A, Bonneau 

M. 2011. Assisted migration: introduction to a multifaceted 

concept. The Forestry Chronicle 87:724-730.

Type: Journal 

Geographic Area: Canada

Keywords: assisted migration, Canada’s forests, climate 

change, forest management, adaptation, species migration

Abstract: The idea that humans can assist nature by purposely 

moving species to suitable habitats to fill the gap between their 

migration capability and the expected rate of climate change 

is being increasingly contemplated and debated as an adaptive 

management option. The interest in assisted migration, both 
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 135

CHAPTER 3 • MIGRATION • GENERAL

Keywords: assisted migration, strategies, review, forest 

management

Abstract: Studies show that changes in climate may exceed 

plant adaptation and migration. The mismatch in rates between 

climate change and plant adaptation and migration will pose 

significant challenges for practitioners that select, grow, and 

outplant native tree species. Native tree species and populations 

that are planted today must meet the climatic challenges that 

they will face during this century. Anticipated shifts in climate 

call for the revision of ethical, legal, political, and economical 

paradigms, as well as changes in the guidelines for growing 

and outplanting trees to maximize survival and curtail maladaptation. Growing trees that survive may be more important 

than growing perfectly shaped trees and may require selection 

of adapted genetic material and/or movement of plant populations (for example, assisted migration). We review and explore 

assisted migration as an adaptation strategy, present some 

working examples, and provide resources for consultation.

Link: http://www.treesearch.fs.fed.us/pubs/46582

378. Watts A. 2014. Assisted migration: growing forests 

adapted to the future. The Forestry Source 19:1, 3.

Type: Newsletter

Geographic Area: North America

Compilers’ Keywords: forest management, silviculture, 

Assisted Migration Adaptation Trial, Society of American 

Foresters

Introduction: Of the management strategies available to create resilient, productive forests in a changing climate, assisted 

migration is one strategy that is prompting much discussion 

in the forestry community. Assisted migration is viewed as a 

proactive strategy because many tree species do not have the 

ability to adapt or migrate naturally at the same rate as the climate is expected to change. This mismatch between trees and 

their environment could result in forests that are less productive and unhealthy.

379. Williams MI, Dumroese RK. 2013. Growing 

assisted migration: synthesis of a climate change adaptation 

strategy. In: Haase DL, Pinto JR, Wilkinson KM, editors. 

National Proceedings: Forest and Conservation Nursery 

Associations—2012. Fort Collins (CO): USDA Forest Service, 

Rocky Mountain Research Station. Proceedings RMRS-P-69. 

p 90-96.

Type: Government Document

Geographic Area: North America

Keywords: climate change, decision framework, implementation, managed relocation, native plant transfer guidelines, 

seed transfer zones

Abstract: Assisted migration may be necessary as a climate 

change adaptation strategy for native plant species that are less 

adaptive or mobile. Moving plants has been practiced a long 

time in human history, but movement of species in response 

to climate change is a new context. First proposed in 1985, 

assisted migration has gained attention since 2007 as a strategy 

to prevent species extinction, minimize economic loss, and 

sustain ecosystem services. We present a synthesis of proposed 

assisted migration guidelines and provide resources for nurseries, landowners, and researchers.

Link: http://www.treesearch.fs.fed.us/pubs/45634

380. Williams MI, Dumroese RK. 2014. Planning the 

future’s forests with assisted migration. In: Sample VA, Bixler 

RP, editors. Forest Conservation and Management in the 

Anthropocene: Adaptation of Science, Policy, and Practices. 

Fort Collins (CO): USDA Forest Service, Rocky Mountain 

Research Station. Proceedings RMRS-P-71. p 133-144. 

Type: Government Document

Geographic Area: Global
136 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

CHAPTER 3 • MIGRATION • RESEARCH

381. Ager A, Nordh NE, Ledin S, Ostry M, Carlson 

M, Ronnberg-Wastljung A. 1990. International transfer of Alnus, Populus, and Salix germplasm: early test results.

Biomass 22:49-62.

Type: Journal 

Geographic Area: Canada, USA, Sweden 

Keywords: biomass, alders, poplars, willows, energy forestry, 

tree improvement

Abstract: The interim results of a genetic testing program 

established within the International Energy Agency (lEA) 

Forestry Energy Agreement are reported. The ‘Joint Evaluation 

Activity’ was initiated to evaluate Alnus, Populus and Salix

germplasm collected by lEA projects and national programs. 

The project was designed to compare the growth rates of selected genetic materials under a variety of growing environments, 

and identify germplasm with potential for biomass cultivation. 

The project was also designed to examine the limitations of 

long-distance international transfer of alder, poplar, and willow clones/provenances ~ species among national biomass 

programs. Four field tests were established, each containing 

selected clones and provenances of alder poplar and willow. 

Three-year growth data are reported here that identify genetic 

materials with superior growth potential in one or more of the 

test environments. They also identify the maximum climatic 

tolerance of several previously untested species/provenances. 

The results can be used to guide the selection of materials for 

future exchanges among the national programs participating in 

the study.

382. Aitken SN, Whitlock MC. 2013. Assisted gene 

flow to facilitate local adaptation to climate change. Annual 

Review of Ecology, Evolution, and Systematics 44:367-388.

Type: Journal 

Geographic Area: Global

Keywords: assisted migration, evolutionary rescue, genetic rescue, outbreeding depression, phenotypic plasticity, 

seed transfer, ecological restoration, reforestation, facilitated 

adaptation

Abstract: Assisted gene flow (AGF) between populations has 

the potential to mitigate maladaptation due to climate change. 

However, AGF may cause outbreeding depression (especially 

if source and recipient populations have been long isolated) 

and may disrupt local adaptation to nonclimatic factors. 

Selection should eliminate extrinsic outbreeding depression 

due to adaptive differences in large populations, and simulations suggest that, within a few generations, evolution should 

resolve mild intrinsic outbreeding depression due to epistasis. 

General (339–380)—debates, reviews, 

trends, surveys, summaries

Research (381–540)—migration studies, 

range shifts, reintroductions

Strategies (541–580)—adaptation options, 

frameworks, decision support

Resources (581–582)—tools, websites, 

software

Migration

#. Author(s), Date. Title. Source.

Type: 

Geographic Area: 

Keywords: 

Description:

Link:

location of study

keywords written by the 

author(s) or provided by the 

compilers

ID number and

information

abstract, summary, or 

portions of the introduction 

or conclusion written by the 

author(s) or provided by the 

compilers

book, book section, conference 

paper, dissertation, government 

document, journal article, 

newsletter, presentation, report, 

thesis or web page

internet address of reference

Each reference contains the following:
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 137

CHAPTER 3 • MIGRATION • RESEARCH

Geographic Area: Northern Hemisphere

Keywords: adaptive traits, conifers, local adaptation, natural 

selection, phenotypic plasticity, provenance trial, quantitative 

genetics

Abstract: Evolutionary responses are required for tree populations to be able to track climate change. Results of 250 years 

of common garden experiments show that most forest trees 

have evolved local adaptation, as evidenced by the adaptive 

differentiation of populations in quantitative traits, reflecting 

environmental conditions of population origins. On the basis 

of the patterns of quantitative variation for 19 adaptation-related traits studied in 59 tree species (mostly temperate and 

boreal species from the Northern Hemisphere), we found that 

genetic differentiation between populations and clinal variation along environmental gradients were very common (respectively, 90% and 78% of cases). Thus, responding to climate change will likely require that the quantitative traits of 

populations again match their environments. We examine what 

kind of information is needed for evaluating the potential to 

respond, and what information is already available. We review 

the genetic models related to selection responses, and what is 

known currently about the genetic basis of the traits. We address special problems to be found at the range margins, and 

highlight the need for more modeling to understand specific issues at southern and northern margins. We need new common 

garden experiments for less known species. For extensively 

studied species, new experiments are needed outside the current ranges. Improving genomic information will allow better 

prediction of responses. Competitive and other interactions 

within species and interactions between species deserve more 

consideration. Despite the long generation times, the strong 

background in quantitative genetics and growing genomic 

resources make forest trees useful species for climate change 

research. The greatest adaptive response is expected when 

populations are large, have high genetic variability, selection is 

strong, and there is ecological opportunity for establishment of 

better adapted genotypes.

Link: http://www.ncbi.nlm.nih.gov/pubmed/23505261

385. Albrecht MA, Guerrant EO Jr, Maschinski J, 

Kennedy KL. 2011. A long-term view of rare plant reintroduction. Biological Conservation 144:2557-2558.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: rare plant conservation, literature 

review

Introduction: Drawing on a literature review and survey 

questionnaires, Godefroid et al. (2011) explore how demographic, genetic, and ecological factors influence success rates 

in plant reintroductions and present valuable recommendations 

to improve plant reintroduction success. But we are concerned 

that the generally dismal picture they paint may erroneously be 

To weigh the risks of AGF against those of maladaptation due 

to climate change, we need to know the species’ extent of local adaptation to climate and other environmental factors, as 

well as its pattern of gene flow. AGF should be a powerful tool 

for managing foundation and resource-producing species with 

large populations and broad ranges that show signs of historical adaptation to local climatic conditions.

Link: http://www.annualreviews.org/doi/abs/10.1146/annurev-ecolsys-110512-135747

383. Aitken SN, Yeaman S, Holliday JA, Wang T, 

Curtis-McLane S. 2008. Adaptation, migration or extirpation: climate change outcomes for tree populations. Evolutionary 

Applications 1:95-111.

Type: Journal 

Geographic Area: Global

Keywords: conifer, ecological genetics, forest, gene flow, genomics, population, selection, species distribution models

Abstract: Species distribution models predict a wholesale redistribution of trees in the next century, yet migratory responses necessary to spatially track climates far exceed maximum 

post-glacial rates. The extent to which populations will adapt 

will depend upon phenotypic variation, strength of selection, 

fecundity, interspecific competition, and biotic interactions. 

Populations of temperate and boreal trees show moderate to 

strong clines in phenology and growth along temperature gradients, indicating substantial local adaptation. Traits involved 

in local adaptation appear to be the product of small effects 

of many genes, and the resulting genotypic redundancy combined with high fecundity may facilitate rapid local adaptation 

despite high gene flow. Gene flow with preadapted alleles from 

warmer climates may promote adaptation and migration at the 

leading edge, while populations at the rear will likely face 

extirpation. Widespread species with large populations and 

high fecundity are likely to persist and adapt, but will likely 

suffer adaptation lag for a few generations. As all tree species 

will be suffering lags, interspecific competition may weaken, 

facilitating persistence under suboptimal conditions. Species 

with small populations, fragmented ranges, low fecundity, or 

suffering declines due to introduced insects or diseases should 

be candidates for facilitated migration.

Link: http://www3.botany.ubc.ca/rieseberglab/plantevol/Aitken_et_al_2008.pdf

384. Alberto FJ, Aitken SN, Alia R, GonzalezMartinez SC, Hanninen H, Kremer A, Lefevre F, 

Lenormand T, Yeaman S, Whetten R, Savolainen 

O. 2013. Potential for evolutionary responses to climate 

change—evidence from tree populations. Global Change 

Biology 19:1645-1661.

Type: Journal 
138 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

CHAPTER 3 • MIGRATION • RESEARCH

Type: Web Page

Geographic Area: Western North America

Compilers’ Keywords: forest management, climate change

Summary: Approximately 200 million seedlings are planted in BC each year. When those trees are harvested 60-80 

years after they are planted, the climate could be 3-4 degrees 

warmer than when the seedlings were planted, exposing 

the trees to maladaptation and health risks. Consequently, 

Ministry of Forests, Land and Natural Resource Operations 

researchers have initiated a large, long-term climate change 

research study—the Assisted Migration Adaptation Trial 

(AMAT)—to better understand tree species’ climate tolerances. Seeds from 15 species growing in BC and neighboring U.S. states were planted between 2009 and 2012 at 48 reforestation sites from northern California to southern Yukon. 

Their growth and health will be monitored, and related to the 

climate of the plantations, enabling researchers to identify 

the seed sources most likely to be best adapted to current 

and future climates. The information will be used to revise 

BC’s species and seed source selection guidelines, helping 

to ensure maximum health and productivity of BC’s planted 

forests well into the future.

Link: http://www.for.gov.bc.ca/hre/forgen/interior/AMAT.

htm

388. Beardmore T, Winder R. 2011. Review of sciencebased assessments of species vulnerability: contributions to 

decision-making for assisted migration. The Forestry Chronicle 

87:745-754.

Type: Journal 

Geographic Area: Canada

Keywords: climate change, forest species, vulnerability 

assessments

Abstract: Assessing climate change impacts on forest species can significantly assist forest management planning. 

Recently, many tools have been developed for assessing 

species-specific vulnerability to climate change. These tools 

are question-based assessments that consider multiple criteria for individual species; the criteria are related to exposure 

and sensitivity to climate change. The following tools are discussed in relation to their use in Canada: (1) the NatureServe 

Climate Change Vulnerability Index; (2) the System for 

Assessing Vulnerability of Species to Climate Change 

(SAVS); (3) the Forest Tree Genetic Risk Assessment; (4) 

the Index for Predicting Tree Species Vulnerability; (5) 

ecological standards developed for the assisted migration 

of Torreya taxifolia; and (6) the Seeds of Success Program. 

These tools can all be applied to different forest species and 

they vary in such areas as their species-specific evaluation 

criteria, means for addressing uncertainty, and the integration of climate change models.

viewed by conservation practitioners, land managers, and policy makers as being broadly representative of reintroduction in 

general, causing hesitation or even dismissing reintroduction 

as a beneficial conservation tool.

386. Anderson AS, Reside AE, VanDerWal JJ, Shoo 

LP, Pearson RG, Williams SE. 2012. Immigrants and 

refugees: the importance of dispersal in mediating biotic attrition 

under climate change. Global Change Biology 18:2126-2134.

Type: Journal 

Geographic Area: Australia

Keywords: climate change, global warming, MAXENT, 

montane tropics, ornithology, range shift

Abstract: Montane tropical rainforests are critically important 

areas for global bird diversity, but are projected to be highly 

vulnerable to contemporary climate change. Upslope shifts of 

lowland species may partially offset declines in upland species but also result in a process of lowland biotic attrition. This 

latter process is contingent on the absence of species adapted 

to novel warm climates, and isolation from pools of potential 

colonizers. In the Australian Wet Tropics, species distribution 

modelling has forecast critical declines in suitable environmental areas for upland endemic birds, raising the question of 

the future role of both natural and assisted dispersal in species 

survival, but information is lacking for important neighbouring 

rainforest regions. Here we use expanded geographic coverage 

of data to model the realized distributions of 120 bird species 

found in north-eastern Australian rainforest, including species 

from potential source locations in the north and recipient locations in the south. We reaffirm previous conclusions as to 

the high vulnerability of this fauna to global warming, and 

extend the list of species whose suitable environmental area 

is projected to decrease. However, we find that expansion of 

suitable area for some species currently restricted to northern 

rainforests has the potential to offset biotic attrition in lowland 

forest of the Australian Wet Tropics. By examining contrasting 

dispersal scenarios, we show that responses to climate change 

in this region may critically depend on dispersal limitation, as 

climate change shifts the suitable environmental envelopes 

of many species south into currently unsuitable habitats. For 

lowland and northern species, future change in vegetation connectivity across contemporary habitat barriers is likely to be 

an important mediator of climate change impacts. In contrast, 

upland species are projected to become increasingly isolated 

and restricted. Their survival is likely to be more dependent 

on the viability of assisted migration, and the emergence and 

persistence of suitable environments at recipient locations.

387. BC Ministry of Forests. 2009. Assisted migration adaptation trial. British Columbia, Canada: BC Ministry 

of Forests, Land and Natural Resource Operations. (URL accessed 29 November 2015)
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 139

CHAPTER 3 • MIGRATION • RESEARCH

plant whose seeds are well dispersed. We reciprocally transplanted seedlings among several populations of different size 

within and among three European regions (in the northwest 

Czech Republic, central Germany and the central Netherlands) 

and studied several fitness-related traits over two growing 

seasons. The region in which the reciprocal transplant experiment was carried out had no influence on the performance of 

seedlings, indicating that there were no differences in overall 

habitat quality. In contrast, the site within region, and the plot 

within site strongly influenced mean plant performance. Plants 

from different populations of origin differed in their performance, indicating genetic variation among populations, but 

performance strongly depended on the specific combination 

of population of origin and transplant site. Plants that grew at 

their home site produced on average almost twice the number 

of seeds per transplant (a multiplicative fitness measure) than 

foreign plants originating from other sites. Survival, rosette 

size and multiplicative fitness all decreased with increasing 

distance from the home site to the transplant site. The size of 

the population of origin did not influence overall plant performance or the strength of local adaptation. In conclusion, our 

results indicate that the common and widespread H. radicata 

consists of locally adapted genotypes within its European range 

at a relatively small scale. Thus a large potential for gene flow 

by seeds and a high density of populations do not appear to 

be sufficient to prevent population differentiation by selection.

391. Bennie J, Hodgson JA, Lawson CR, Holloway 

CT, Roy DB, Brereton T, Thomas CD, Wilson RJ. 

2013. Range expansion through fragmented landscapes under 

a variable climate. Ecology Letters 16:921-929.

Type: Journal 

Geographic Area: United Kingdom

Keywords: climate variability, colonization, environmental 

threshold model, extinction, habitat networks, mechanistic model, metapopulation dynamics, microrefugia, species 

distribution

Abstract: Ecological responses to climate change may depend on complex patterns of variability in weather and local 

microclimate that overlay global increases in mean temperature. Here, we show that high-resolution temporal and spatial 

variability in temperature drives the dynamics of range expansion for an exemplar species, the butterfly Hesperia comma. 

Using fine-resolution (5 m) models of vegetation surface microclimate, we estimate the thermal suitability of 906 habitat 

patches at the species’ range margin for 27 years. Population 

and metapopulation models that incorporate this dynamic microclimate surface improve predictions of observed annual 

changes to population density and patch occupancy dynamics during the species’ range expansion from 1982 to 2009. 

Our findings reveal how fine-scale, short-term environmental 

variability drives rates and patterns of range expansion through 

spatially localized, intermittent episodes of expansion and 

389. Beckage B, Osborne B, Gavin DG, Pucko C, 

Siccama T, Perkins T. 2008. A rapid upward shift of a forest ecotone during 40 years of warming in the Green Mountains 

of Vermont. Proceedings of the National Academy of Sciences 

USA 105:4197-4202.

Type: Journal 

Geographic Area: Northeastern USA

Keywords: climate change, range shifts

Abstract: Detecting latitudinal range shifts of forest trees in 

response to recent climate change is difficult because of slow 

demographic rates and limited dispersal but may be facilitated 

by spatially compressed climatic zones along elevation gradients in montane environments. We resurveyed forest plots 

established in 1964 along elevation transects in the Green 

Mountains (Vermont) to examine whether a shift had occurred 

in the location of the northern hardwood-boreal forest ecotone (NBE) from 1964 to 2004. We found a 19% increase in 

dominance of northern hardwoods from 70% in 1964 to 89% 

in 2004 in the lower half of the NBE. This shift was driven by 

a decrease (up to 76%) in boreal and increase (up to 16%) in 

northern hardwood basal area within the lower portions of the 

ecotone. We used aerial photographs and satellite imagery to 

estimate a 91- to 119-m upslope shift in the upper limits of the 

NBE from 1962 to 2005. The upward shift is consistent with 

regional climatic change during the same period; interpolating 

climate data to the NBE showed a 1.1 °C increase in annual 

temperature, which would predict a 208-m upslope movement 

of the ecotone, along with a 34% increase in precipitation. 

The rapid upward movement of the NBE indicates little inertia to climatically induced range shifts in montane forests; 

the upslope shift may have been accelerated by high turnover 

in canopy trees that provided opportunities for ingrowth of 

lower elevation species. Our results indicate that high-elevation forests may be jeopardized by climate change sooner than 

anticipated.

Link: http://www.ncbi.nlm.nih.gov/pubmed/18334647

390. Becker U, Dostal P, Jorritsma-Wienk LD, 

Matthies D. 2008. The spatial scale of adaptive population 

differentiation in a wide-spread, well-dispersed plant species. 

Oikos 117:1865-1873.

Type: Journal

Geographic Area: Europe

Compilers’ Keywords: adaptation, Hypochoeris radicata, reciprocal transplant, genetic variation, gene flow

Abstract: Adaptation to the specific conditions at different 

sites may contribute strongly to the wide distribution of a plant 

species. However, little is known about the scale at which such 

adaptation occurs in common species. We studied population 

differentiation, plasticity and local adaptation of the short-lived 

perennial Hypochoeris radicata, a widespread and common 
140 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

CHAPTER 3 • MIGRATION • RESEARCH

Compilers’ Keywords: assisted migration, translocation, 

maladaptation, commentary, Laccaria, editorial

Introduction: Plants are being moved across the globe at an 

increasing rate both within and outside their current distributional range. Research in the past few decades has focused intensely on invasive exotic plant species that are moved outside 

their distributional range. Such invaders can have dramatic 

impacts on indigenous communities through altered biotic interactions with competitors, mutualists, and antagonists. But 

likewise, translocation within the distributional range exposes 

nonnative plant genotypes to interactions with nonlocal coevolved competitors, mutualists, and antagonists. A major concern in restoration ecology is that translocation of seeds from 

foreign seed sources introduces plant genotypes that are maladapted to local conditions and subsequently hybridize with 

locally adapted conspecifics, decreasing mean population fitness. Interestingly, while local adaptation of plants to their abiotic environment is well documented, warranting the concern, 

studies of plant local adaptation to their biotic environment 

have shown less unequivocal results, ranging from local adaption to local maladaptation for interactions with antagonists. A 

large reciprocal transplant field study by Cremieux et al. reported in this issue of New Phytologist, offers a prime example 

of the diversity of outcomes of altered biotic interactions that 

we can observe following translocation of plants within their 

current distributional range even within a single community. 

In one plant species, local plant genotypes were more resistant 

to the local demes of specialist antagonist than nonlocal plant 

genotypes, suggesting plant local adaptation to important aspects of the biotic environment. However, in another plant species, the reverse pattern was observed. The take-home message 

from this study is a teeth-grinder for restoration ecologists: it 

is hard to make a general prediction as to whether foreign seed 

provenances pose a risk of introducing alleles causing low biotic resistance in the restoration area or not. 

394. Bodin J, Badeau V, Bruno E, Cluzeau C, 

Moisselin J-M, Walther GR, Dupouey J-L. 2013.

Shifts of forest species along an elevational gradient in Southeast 

France: climate change or stand maturation? Journal of 

Vegetation Science 24:269-283.

Type: Journal 

Geographic Area: Southeast France

Keywords: elevation, forest dynamics, forest inventory, landuse change, long-term monitoring, mountains of Southeast 

France, niche, modelling, plant species shift

Abstract: Recent vegetation changes in mountain areas are 

often explained by climate warming. However, effects of landuse changes, such as recolonization of abandoned pastures by 

forest, are difficult to separate from those of climate change. 

Even within forest belts, changes in stand structure due to 

forest management and stand maturation could confound the 

contraction. Incorporating dynamic microclimates can thus 

improve models of species range shifts at spatial and temporal 

scales relevant to conservation interventions.

Link: http://www.ncbi.nlm.nih.gov/pubmed/23701124

392. Bennuah SY, Wang T, Aitken SN. 2004.

Genetic analysis of the Picea sitchensis x glauca introgression 

zone in British Columbia. Forest Ecology and Management 

197:65-77.

Type: Journal 

Geographic Area: Western Canada

Keywords: Sitka spruce, white spruce, introgression zone, 

hybrid index, STS markers, allozymes, geographic pattern

Abstract: A genetic analysis of the introgression zone between Sitka spruce (Picea sitchensis) and white spruce (P. 

glauca) in northwestern British Columbia was conducted 

to (1) quantify the contributions of parental species to introgressed populations; (2) characterize genetic patterns 

of variation across the introgression zone; and (3) address 

practical issues of gene resource management in this region. 

Seeds were sampled and seedlings grown from 57 putative 

hybrid parent trees from 16 locations across the introgression zone. Foliage samples were also obtained for allopatric 

Sitka and white spruce reference populations. Based on allele frequencies for sequence-tagged-site (STS) molecular 

markers, a hybrid index was developed that reflects the 

relative contribution of Sitka spruce to hybrid populations (0 

= white spruce, 1 = Sitka spruce). Mean STS hybrid index 

estimates ranged from 0.46 to 0.95 for locations within the 

zone of introgression. Significant differences in hybrid index 

were observed among sampling locations. Hybrid indices 

decreased from the coast to the interior within the introgression zone in spite of large among family variation within 

locations. Variation among locations in hybrid index was 

better explained by geographic variables than by climatic 

or ecological variables. Physical drainage distance along 

river valleys to the nearest marine inlet explained 78% of 

the variation in STS hybrid index. Concordant clines were 

obtained when the same analytical techniques were applied 

to published allozyme frequencies for seedlots originating 

from a wider geographic area across the introgression zone. 

Allozyme hybrid index estimates ranged from 0.01 to 0.98. 

The relatively steep cline observed in hybrid index across 

the maritime/continental climate ecotone is best managed by 

limiting longitudinal seed transfer for reforestation in this 

region.

393. Biere A, Verhoeven KJF. 2008. Local adaptation 

and the consequences of being dislocated for coevolved enemies. 

New Phytologist 180:265-268.

Type: Journal 

Geographic Area: Global
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 141

predominantly in winter or with a 20% decrease in annual 

precipitation, or (2) if 2 °C warming occurs, with or without 

decrease in annual precipitation, and regardless of whether 

there is a uniform monthly warming pattern or one that is 

higher in winter than in summer. Available information concerning other factors that might also affect climatic-change 

responses suggests that these large predicted impacts on temperate Florida species may be underestimates. Subtropical 

Florida species will tend to move north and inland with warming but extensive human assistance may be needed, if they are 

to realize their newly expanded, potential natural ranges.

396. Bradley BA, Blumenthal DM, Early R, Grosholz 

ED, Lawler JJ, Miller LP, Sorte CJB, D’Antonio CM, 

Diez JM, Dukes JS, Ibanez I, Olden JD. 2012. Global 

change, global trade, and the next wave of plant invasions.

Frontiers in Ecology and the Environment 10:20-28.

Type: Journal 

Geographic Area: North America

Compilers’ Keywords: non-native plants, climate change, 

horticulture trade, drought-tolerance, migration, review

Abstract: Many non-native plants in the U.S. have become 

problematic invaders of native and managed ecosystems, but 

a new generation of invasive species may be at our doorstep. 

Here, we review trends in the horticultural trade and invasion 

patterns of previously introduced species and show that novel 

species introductions from emerging horticultural trade partners are likely to rapidly increase invasion risk. At the same 

time, climate change and water restrictions are increasing demand for new types of species adapted to warm and dry environments. This confluence of forces could expose the U.S. to 

a range of new invasive species, including many from tropical 

and semiarid Africa as well as the Middle East. Risk assessment strategies have proven successful elsewhere at identifying and preventing invasions, although some modifications are 

needed to address emerging threats. Now is the time to implement horticulture import screening measures to prevent this 

new wave of plant invasions.

Link: http://web.ics.purdue.edu/~jsdukes/BradleyEtAl2012.

pdf

397. Brooker RW, Travis JM, Clark EJ, Dytham C. 

2007. Modelling species’ range shifts in a changing climate: the 

impacts of biotic interactions, dispersal distance and the rate of 

climate change. Journal of Theoretical Biology 245:59-65.

Type: Journal 

Geographic Area: Global

Keywords: climate change, biodiversity, range shifting, biotic 

interactions, dispersal, modelling

Abstract: There is an urgent need for accurate prediction of 

climate change impacts on species ranges. Current reliance on 

climate signal. Here, we evaluate the direction and rate of 

plant species elevation shifts in mountain forests, considering 

the role of stand dynamics. We compared floristic data from 

the French National Forest Inventory collected in the 1980s 

and 1990s. They provided a large-scale (30,985 plots) and representative sample of vegetation between 0 and 2500 m a.s.l. 

Species response curves along the elevation and exposure gradients were fitted with a logistic regression model. In order to 

assess the effect of changes in successional stages of the forest 

stands, we compared plant species shifts in the whole set of 

stands with those solely in closed stands. A total of 62 species 

shifted downward, whereas 113 shifted upward, resulting in a 

significant upward mean shift of 17.9 m. Upward shifting species were preferentially woody and heliophilous, suggesting a 

role for forest closure and maturation in the observed changes. 

Excluding all open forest stages from analyses, the upward 

trend became weaker (-3.0 m) and was not significant. Forests 

of the study area have undergone closure and maturation, more 

strongly at lower altitudes than at higher ones, producing an 

apparent shift of species. In the mountain relief of Southeast 

France, changes in the successional stages of stands appear 

as the main cause of the apparent upslope movement of forest species. Since a similar trend of forest maturation exists in 

large areas throughout Europe, forest dynamics should be better taken into account among the causes of vegetation changes 

before inferring any climate change effect.

Link: http://onlinelibrary.wiley.com/doi/10.1111/j.1654-

1103.2012.01456.x/full

395. Box EO, Crumpacker DW, Hardin ED. 1999.

Predicted effects of climate change on distribution of ecologically important native tree and shrub species in Florida. Climatic 

Change 41:213-248.

Type: Journal 

Geographic Area: Florida, USA

Compilers’ Keywords: climatic envelope model, temperate 

species, tropical species

Abstract: A previously developed plant species-climatic envelope model was evaluated further and used to predict effects 

of hypothesized climatic change on the potential distribution 

of 124 native woody plant species in Florida, U.S.A. Twelve 

scenarios were investigated. These included mean annual temperature increases of 1 °C or 2 °C, achieved either by equal 1 

°C or 2 °C increases on a monthly basis throughout the year, or 

by disproportionately larger seasonal increases in winter and 

smaller ones in summer. The various temperature increases 

were then combined with each of several precipitation changes, ranging from +10% to –20%, to produce the final set of 

scenarios. More detailed analysis involving six of the scenarios 

and a subset of 28 representative, ecologically important species suggested that (1) large decreases in the Florida range of 

many temperate species would result if 1 °C warming occurs 

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in major genebanks—promising locations in which to focus 

future genetic resource conservation efforts.

Link: http://www.stanford.edu/~mburke/papers/burke%20

et%20al%20GEC%202009.pdf

399. Burrows MT, Schoeman DS, Richardson 

AJ, Molinos JG, Hoffmann A, Buckley LB, Moore 

PJ, Brown CJ, Bruno JF, Duarte CM, Halpern BS, 

Hoegh-Guldberg O, Kappel CV, Kiessling W, 

O’Connor MI, Pandolfi JM, Parmesan C, Sydeman 

WJ, Ferrier S, Williams KJ, Poloczanska ES. 2014. 

Geographical limits to species-range shifts are suggested by climate velocity. Nature 507:492-495.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: migration, biodiversity, climate 

change, climate niche, species distributions, modelling, 

trajectories

Abstract: The reorganization of patterns of species diversity driven by anthropogenic climate change, and the consequences for humans, are not yet fully understood or appreciated. Nevertheless, changes in climate conditions are 

useful for predicting shifts in species distributions at global 

and local scales. Here we use the velocity of climate change 

to derive spatial trajectories for climatic niches from 1960 

to 2009 (ref. 7) and from 2006 to 2100, and use the properties of these trajectories to infer changes in species distributions. Coastlines act as barriers and locally cooler areas act 

as attractors for trajectories, creating source and sink areas 

for local climatic conditions. Climate source areas indicate 

where locally novel conditions are not connected to areas 

where similar climates previously occurred, and are thereby 

inaccessible to climate migrants tracking isotherms: 16% of 

global surface area for 1960 to 2009, and 34% of ocean for 

the ‘business as usual’ climate scenario (representative concentration pathway (RCP) 8.5) representing continued use of 

fossil fuels without mitigation. Climate sink areas are where 

climate conditions locally disappear, potentially blocking the 

movement of climate migrants. Sink areas comprise 1.0% of 

ocean area and 3.6% of land and are prevalent on coasts and 

high ground. Using this approach to infer shifts in species 

distributions gives global and regional maps of the expected 

direction and rate of shifts of climate migrants, and suggests 

areas of potential loss of species richness.

Link: http://www.ncbi.nlm.nih.gov/pubmed/24509712

400. Byrne M, Stone L, Millar MA. 2011. Assessing 

genetic risk in revegetation. Journal of Applied Ecology 

48:1365-1373.

Type: Journal 

Geographic Area: Global

bioclimatic envelope approaches ignores important biological 

processes such as interactions and dispersal. Although much 

debated, it is unclear how such processes might influence 

range shifting. Using individual-based modelling we show that 

interspecific interactions and dispersal ability interact with the 

rate of climate change to determine range-shifting dynamics 

in a simulated community with two growth forms—mutualists 

and competitors. Interactions determine spatial arrangements 

of species prior to the onset of rapid climate change. These 

lead to space-occupancy effects that limit the rate of expansion 

of the fast-growing competitors but which can be overcome 

by increased long-distance dispersal. As the rate of climate 

change increases, lower levels of long-distance dispersal can 

drive the mutualists to extinction, demonstrating the potential 

for subtle process balances, non-linear dynamics and abrupt 

changes from species coexistence to species loss during climate change.

Link: http://www.ncbi.nlm.nih.gov/pubmed/17087974

398. Burke MB, Lobell DB, Guarino L. 2009.

Shifts in African crop climates by 2050, and the implications for 

crop improvement and genetic resources conservation. Global 

Environmental Change 19:317-325.

Type: Journal 

Geographic Area: Africa

Keywords: maize, Africa, genetic resources, climate change, 

adaptation, sorghum, millet

Abstract: Increased understanding of the substantial threat 

climate change poses to agriculture has not been met with a 

similarly improved understanding of how best to respond. 

Here we examine likely shifts in crop climates in Sub-Saharan 

Africa under climate change to 2050, and explore the implications for agricultural adaptation, with particular focus on 

identifying priorities in crop breeding and the conservation 

of crop genetic resources. We find that for three of Africa’s 

primary cereal crops—maize, millet, and sorghum—expected 

changes in growing season temperature are considerable and 

dwarf changes projected for precipitation, with the warmest recent temperatures on average cooler than almost 9 out 

of 10 expected observations by 2050. For the ‘‘novel’’ crop 

climates currently unrepresented in each country but likely 

extant there in 2050, we identify current analogs across the 

continent. The majority of African countries will have novel 

climates over at least half of their current crop area by 2050. 

Of these countries, 75% will have novel climates with analogs 

in the current climate of at least five other countries, suggesting that international movement of germplasm will be necessary for adaptation. A more troubling set of countries—largely 

the hotter Sahelian countries—will have climates with few 

analogs for any crop. Finally, we identify countries, such as 

Sudan, Cameroon, and Nigeria, whose current crop areas are 

analogs to many future climates but that are poorly represented 

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completion of the annual cycle of the species and therefore on 

their accomplished phenology. In particular, they have shown 

that the northern limit of species’ ranges appears to be caused 

mainly by the inability to undergo full fruit maturation, while 

the southern limit appears to be caused by the inability to flower or unfold leaves owing to a lack of chilling temperatures 

that are necessary to break bud dormancy. I discuss here why 

phenology is a key adaptive trait in shaping species distribution using mostly examples from plant species, which have 

been the most documented. After discussing how phenology is 

involved in fitness and why it is an adaptive trait susceptible to 

evolve quickly in changing climate conditions, I describe how 

phenology is related to fitness in species distribution processbased models and discuss the fate of species under climate 

change scenarios using model projections and experimental or 

field studies from the literature.

Link: http://www.ncbi.nlm.nih.gov/pubmed/20819809

402. Corlett RT, Westcott DA. 2013. Will plant movements keep up with climate change? Trends in Ecology and 

Evolution 28:482-488.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: review, migration, velocity, tracking

Abstract: In the face of anthropogenic climate change, species must acclimate, adapt, move, or die. Although some species are moving already, their ability to keep up with the faster 

changes expected in the future is unclear. ‘Migration lag’ is a 

particular concern with plants, because it could threaten both 

biodiversity and carbon storage. Plant movements are not realistically represented in models currently used to predict future 

vegetation and carbon-cycle feedbacks, so there is an urgent 

need to understand how much of a problem failure to track 

climate change is likely to be. Therefore, in this review, we 

compare how fast plants need to move with how fast they can 

move; that is, the velocity of climate change with the velocity 

of plant movement.

Link: http://www.ncbi.nlm.nih.gov/pubmed/23721732

403. Damschen EI, Harrison S, Ackerly DD, 

Fernández-Going BM, Anacker BL. 2012. Endemic 

plant communities on special soils: early victims or hardy survivors of climate change? Journal of Ecology 100:1122-1130.

Type: Journal 

Geographic Area: Global

Keywords: assisted migration, azonal vegetation, community, conservation, edaphic, functional diversity, managed 

relocation, plant-climate interactions, risk, serpentine, species 

diversity

Abstract: Predicting and mitigating climate change effects 

on ecological communities is a tremendous challenge. Little 

Keywords: gene flow, genetic change, hybridisation, outbreeding depression, pollen dispersal, risk assessment

Abstract: Revegetation can provide major environmental 

benefits in degraded landscapes, but there is also potential for 

negative impacts from genetic change in local native populations. Broad areas of revegetation may provide a large source of 

foreign genes in landscapes where small remnant native populations act as a sink. Genetic change from hybridisation can 

threaten population persistence and contribute to species extinction through genetic assimilation or demographic swamping. Implementation of revegtation within a risk management 

framework allows identification of risk factors, analysis and 

evaluation of risk to inform decision-making and management to minimize and mitigate the risk. Informed analysis and 

evaluation of genetic risk is important in revegetation because 

it will be difficult to control or reverse the impacts in natural 

ecosystems and they are often not expressed until the second 

generation or later. A risk assessment protocol is presented 

based on evaluation of factors that influence the likelihood and 

consequences of adverse genetic change from revegetation 

arising through pollen dispersal. The assessment is applicable 

to a broad ranges of revegetation activities and contributes to 

the development of informed decision-making processes in 

implementation of revegetation systems and land use practices 

that protect and enhance biodiversity in degraded landscapes. 

Implementation of revegetation programmes within a risk 

management framework will help to ensure that significant environmental benefits are captured with minimal concomitant 

negative impacts on the surrounding biodiversity. A genetic 

risk protocol provides a tool for evaluation of potential adverse 

genetic impacts on native populations from revegetation and 

can be implemented in conjunction with weed risk assessment. 

Risk assessment as an integral part of evaluation of environmental impact for large-scale revegetation programmes will 

contribute to the development of informed decision-making 

processes in the implementation of revegetation systems, and 

ultimately, it will aid in the development of land uses that protect and enhance biodiversity in degraded landscapes. 

401. Chuine I. 2010. Why does phenology drive species 

distribution? Philosophical Transactions of the Royal Society 

B: Biological Sciences 365:3149-3160.

Type: Journal 

Geographic Area: Global

Keywords: fitness, adaptation, niche, species distribution 

Abstract: Despite the numerous studies which have been 

conducted during the past decade on species ranges and their 

relationship to the environment, our understanding of how 

environmental conditions shape species distribution is still far 

from complete. Yet, some process-based species distribution 

models have been able to simulate plants and insects distribution at a global scale. These models strongly rely on the 

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expansion in the last c. 500 years. Positive and negative cultural legacies are evident and intervention is considered a necessary restoration tool. Managed grazing at key stages of habitat 

development can support continued recruitment and regeneration. A more flexible and integrated management approach is 

recommended. Long-term ecology can provide direction and 

address uncertainties, while ecological and stakeholder evidence provide the detail necessary to develop effective management that incorporates ecosystem perspectives. This can 

help shape management strategies that balance possible conflicts over perceived short-term ‘damage’ to secure longer term 

processes. Learning based on multiple temporal perspectives 

has the potential to contribute to conservation and restoration 

planning and practice. These principles are more widely applicable in adaptive management.

405. Diaz S, Cabido M. 1997. Plant functional types 

and ecosystem function in relation to global change. Journal of 

Vegetation Science 8:463-474.

Type: Journal 

Geographic Area: Global

Keywords: Argentina, climate change, plant functional trait, 

regional gradient

Abstract: Plant functional types (PFTs) bridge the gap between 

plant physiology and community and ecosystem processes, 

thus providing a powerful tool in climate change research. We 

aimed at identifying PFTs within the flora of central-western 

Argentina, and to explore their possible consequences for ecosystem function. We analyzed 24 vegetative and regenerative 

traits of the 100 most abundant species along a steep climatic 

gradient. Based on plant traits and standard multivariate techniques, we identified eight PFTs. Our results confirmed, over 

a wide range of climatic conditions, the occurrence of broad 

recurrent patterns of association among plant traits reported 

for other floras; namely trade-offs between high investment in 

photosynthesis and growth on the one hand, and preferential 

allocation to storage and defense on the other. Regenerative 

traits were only partially coupled with vegetative traits. Using 

easily-measured plant traits and individual species cover in 63 

sites, we predicted main community ecosystem processes along 

the regional gradient. We hypothesized likely impacts of global 

climatic change on PFTs and ecosystems in situ, and analysed 

their probabilities of migrating in response to changing climatic 

conditions. Finally, we discuss the advantages and limitations 

of this kind of approach in predicting changes in plant distribution and in ecosystem processes over the next century.

Link: http://www.dpi.inpe.br/referata/arq/16_Andre/Diaz_e_

Cabido_1997.pdf

406. Drayton B, Primack RB. 2012. Success rates for 

reintroductions of eight perennial plant species after 15 years. 

Restoration Ecology 20:299-303.

attention has been given to how endemic-rich communities 

on isolated patches of low-nutrient soil (e.g. serpentine) will 

respond to climate change. To address spatial factors (the isolated nature of outcrops), we incorporate habitat patchiness 

into species distribution models under climate change. The 

degree of overlap between current and future suitable habitat 

does not change when patchy habitats are incorporated, probably because serpentine occurs in mountainous regions where 

climatically and edaphically suitable regions geographically 

coincide. The dispersal distances required to move to newly 

suitable habitat are large, however, making successful migration unlikely. To address how non-spatial factors affect the climate change responses of serpentine plant communities (e.g. 

the impacts of nutrient limitation and stress-tolerant functional 

traits), we conduct a literature review. Some studies suggest 

that serpentine communities may be at less risk than ‘normal’ 

soil communities due to their stress-tolerant functional traits, 

but there is also evidence to the contrary. Assessing climate 

change risk for the world’s diverse edaphic floras requires 

determining impacts on both special and ‘normal’ soil communities. Studies are needed that use functional traits, evaluate the role of evolutionary and ecological plasticity, examine 

responses across spatial and temporal scales and assess the efficacy of managed relocation efforts.

404. Davies AL. 2011. Long-term approaches to native 

woodland restoration: palaeoecological and stakeholder perspectives on Atlantic forests of northern Europe. Forest Ecology 

and Management 261:751-763.

Type: Journal 

Geographic Area: Northern Europe

Keywords: long-term ecology, western oakwood, stakeholder 

perspectives, palaeoecology, adaptive management

Abstract: The long timeframes involved in woodland regeneration and adaptation introduce considerable uncertainty into 

management and conservation planning as most ecological 

datasets span only a small part of ecosystem dynamics. This 

is a particular concern in vulnerable habitats, such as Atlantic 

birch-oak woods in north-western Britain, where range edge 

populations are at risk from herbivory and climate change. 

This study combines historical palaeoecology and stakeholder 

observations to assess how multiple perspectives can inform 

existing models, expectations and goals for Atlantic woodland 

management. Long-term evidence for changing woodland 

composition addresses stakeholder uncertainties over the currently restricted distribution of oak, alder and hazel. Oak has 

remained a secondary component of birch-dominated woods 

and habitat definitions based on oak are too narrow to ensure 

sustainability. Birch has survived numerous periods of climatic 

and biotic adversity and shows the strongest positive response 

to historic reductions in grazing pressure. This has led to a 

pulse of regeneration since c. AD 1900, so current restoration 

efforts are taking place within the strongest period of woodland 

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408. Elith J, Leathwick JR. 2009. Species distribution 

models: ecological explanation and prediction across space and 

time. Annual Review of Ecology, Evolution, and Systematics 

40:677-697.

Type: Journal 

Geographic Area: Global

Keywords: climate change, invasions, niche, predict, presence-only, spatial

Abstract: Species distribution models (SDMs) are numerical 

tools that combine observations of species occurrence or abundance with environmental estimates. They are used to gain ecological and evolutionary insights and to predict distributions 

across landscapes, sometimes requiring extrapolation in space 

and time. SDMs are now widely used across terrestrial, freshwater, and marine realms. Differences in methods between disciplines reflect both differences in species mobility and in “established use.” Model realism and robustness is influenced by 

selection of relevant predictors and modeling method, consideration of scale, how the interplay between environmental and 

geographic factors is handled, and the extent of extrapolation. 

Current linkages between SDM practice and ecological theory 

are often weak, hindering progress. Remaining challenges include: improvement of methods for modeling presence-only 

data and for model selection and evaluation; accounting for 

biotic interactions; and assessing model uncertainty.

Link: http://mgel.env.duke.edu/wp-content/workshop-dropbox/Elith_Leathwick_2009.pdf

409. Elith J, Phillips SJ, Hastie T, Dudík M, Chee 

YE, Yates CJ. 2011. A statistical explanation of MaxEnt for 

ecologists. Diversity and Distributions 17:43-57.

Type: Journal 

Geographic Area: Global

Keywords: absence, ecological niche, entropy, machine learning, presence-only, species distribution model

Abstract: MaxEnt is a program for modelling species distributions from presence only species records. This paper is 

written for ecologists and describes the MaxEnt model from 

a statistical perspective, making explicit links between the 

structure of the model, decisions required in producing a 

modelled distribution, and knowledge about the species and 

the data that might affect those decisions. To begin we discuss 

the characteristics of presence-only data, highlighting implications for modelling distributions. We particularly focus on the 

problems of sample bias and lack of information on species 

prevalence. The keystone of the paper is a new statistical explanation of MaxEnt which shows that the model minimizes 

the relative entropy between two probability densities (one 

estimated from the presence data and one, from the landscape) 

defined in covariate space. For many users, this viewpoint is 

likely to be a more accessible way to understand the model 

Type: Journal 

Geographic Area: Eastern USA

Keywords: plant conservation, plant reintroduction, reintroduction methods

Abstract: The creation of new populations of rare and endangered plant species has become well-established as a 

standard technique in conservation and restoration ecology. 

However, much remains unknown about the actual rates of 

success or failure of such reintroductions. Recent research 

suggests that in part this reflects under-reporting of failures. 

In 2000, the authors published a paper reporting rates of success in reintroducing eight perennial plant species into two 

reserves near Boston, MA, in 1994-1995. In 2010, the authors conducted a census of the experimental sites 15 years 

after reintroduction; almost all the populations reported in 

2000 had disappeared. The implications for reintroduction 

methodology, with respect to establishing and reporting both 

successful and unsuccessful experiments are discussed. 

407. Duran KL, Lowrey TK, Parmenter RR, Lewis 

PO. 2005. Genetic diversity in Chihuahuan Desert populations of creosotebush (Zygophyllaceae: Larrea tridentata).

American Journal of Botany 92:722-729.

Type: Journal 

Geographic Area: Mexico, Southwestern USA

Keywords: biogeography, Chihuahuan Desert, creosotebush, 

genetic variation, isozymes, Larrea tridentata, population 

genetics, Zygophyllaceae

Abstract: We examined isozyme variation in the dominant 

Chihuahuan Desert shrub, Larrea tridentata (creosotebush), 

to determine the genetic variation within and among populations, the biogeographic relationships of populations, and the 

potential inbreeding in the species. We surveyed 17 populations consisting of 20 to 50 individuals per population along a 

1600 km north–south transect across the Chihuahuan Desert. 

The southernmost population was near Villa Hidalgo, Mexico, 

and the northernmost near Isleta Pueblo, New Mexico. All 12 

isozyme loci examined were polymorphic (Ht = 0.416), with 

up to nine alleles per locus. Despite high levels of variation, 

we detected moderate inbreeding in L. tridentata populations. Most variation was found within rather than among 

populations (GST = 0.118). Furthermore, recently established populations in the northern limits of the Chihuahuan 

Desert did not show decreased levels of genetic variation 

(Ho = 0.336). A significant correlation was found between pairwise genetic and geographic distances (r = 0.305). Larrea tridentata showed and continues to show a massive range expansion 

into the arid and semi-arid regions of the American Southwest, 

but as shown by the high genetic variation, this expansion took 

place as a wave, rather than a series of founder events.

Link: http://sev.lternet.edu/sites/default/files/sev306_duran_

etal_ajb_2005.pdf

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results confirm the importance of establishing an appropriate 

relational basis in time and space between species and climatic 

layers, providing key operational criteria for selection of species observations fed into SDMs. Our findings highlight the 

importance of identifying weather stations locations used in 

interpolated products, which will allow a characterization of 

some aspects of uncertainty and identification of regions where 

users need to be particularly careful when making a decision 

based on a SDM.

411. Fischer J, Lindenmayer DB. 2000. An assessment of the published results of animal relocations. Biological 

Conservation 96:1-11.

Type: Journal 

Geographic Area: Global

Keywords: translocation, re-introduction, supplementation, 

relocation, re-introduction success

Abstract: We reviewed 180 case studies and a number of 

theoretical papers on animal relocations published in 12 major international scientific journals over the last 20 years. The 

study focused on re-introductions, supplementations and translocations. We did not assess introductions. Re-introductions 

were the most common type of relocation (116/180); three 

quarters of these were conducted for conservation purposes. 

Supplementations (48/180) and translocations (36/180) occurred less frequently, and both were commonly carried out for 

reasons other than conservation. Simple descriptive statistics 

were used to analyze factors influencing relocation success. 

Translocations that aimed to solve human animal conflicts 

generally failed. Re-introduction success was not found to 

have changed over the last two decades, but re-introductions 

appeared to be more successful when the source population 

was wild, a large number of animals was released (n >100), 

and the cause of original decline was removed. More complex 

trends were found for the effect of predation and the use of supportive measures such as provision of food or shelter, or predator control prior to release. The success of 47% of reintroductions was uncertain at the time case studies were published in 

journals. This was partly due to the lack of generally accepted 

and widely applied criteria to assess success. Very few case 

studies (3%) reported the cost of the relocation attempt. We 

conclude that there were three primary aims for animal relocations. These were to solve human-animal conflicts, to restock 

game populations, and conservation. Our extensive review 

of the present literature leads us to conclude that the value of 

animal relocations as a conservation tool could be enhanced 

through (1) more rigorous testing for the appropriateness of 

the approach in a given case, (2) the establishment of widely 

used and generally accepted criteria for judging the success or 

failure of relocations, (3) better monitoring after a relocation, 

(4) better financial accountability, and (5) greater effort to publish the results of relocations, even ones that are unsuccessful.

than previous ones that rely on machine learning concepts. We 

then step through a detailed explanation of MaxEnt describing 

key components (e.g. covariates and features, and definition 

of the landscape extent), the mechanics of model fitting (e.g. 

feature selection, constraints and regularization) and outputs. 

Using case studies for a Banksia species native to south-west 

Australia and a riverine fish, we fit models and interpret them, 

exploring why certain choices affect the result and what this 

means. The fish example illustrates use of the model with vector data for linear river segments rather than raster (gridded) 

data. Appropriate treatments for survey bias, unprojected data, 

locally restricted species, and predicting to environments outside the range of the training data are demonstrated, and new 

capabilities discussed. Online appendices include additional 

details of the model and the mathematical links between previous explanations and this one, example code and data, and 

further information on the case studies.

Link: http://nate2.myweb.uga.edu/FANR8400/Elith_

etal_2010_maxent%20for%20ecologists.pdf

410. Fernández M, Hamilton H, Kueppers LM. 

2013. Characterizing uncertainty in species distribution models derived from interpolated weather station data. Ecosphere 

4:1-17.

Type: Journal 

Geographic Area: North America

Keywords: climate-based niche models, gridded climate, 

uncertainty

Abstract: Species distribution models (SDMs) are used to 

generate hypotheses regarding the potential distributions of 

species under different environmental conditions, such as forecasts of species range shifts in response to climate change and 

predictions of invasive species range expansions. However, an 

accurate description of species’ geographic ranges as a function of the environment requires that species observations 

and climatic variables are measured at the same spatial and 

temporal resolution, which is usually not the case. Weather 

station data are interpolated and these resulting continuous 

data layers are incorporated into SDMs, often without any 

uncertainty assessment. Here we quantify the effects of three 

unrelated but complementary aspects of uncertainty in weather 

station interpolations on SDM performance using MaxEnt. We 

examine the influence of topographic heterogeneity, interannual variability, and distance to station on the over- and under-prediction of modeled North American bird distributions. 

Our species observations are derived from presence-absence 

information for 20 bird species with well-known distributions. 

These three metrics of uncertainty in interpolated weather station data have varying contributions to over- and under prediction errors in SDMs. Topographic heterogeneity had the highest contribution to omission errors; the lowest contribution to 

commission errors was from Euclidean distance to station. The 

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(on average 52%, 19% and 16%, respectively). Furthermore, 

our results show a success rate decline in individual experiments with time. Survival rates reported in the literature are 

also much higher (78% on average) than those mentioned 

by survey participants (33% on average). We identified various parameters that positively influence plant reintroduction 

outcomes, e.g., working in protected sites, using seedlings, 

increasing the number of reintroduced individuals, mixing material from diverse populations, using transplants from stable 

source populations, site preparation or management effort and 

knowledge of the genetic variation of the target species. This 

study also revealed shortcomings of common experimental designs that greatly limit the interpretation of plant reintroduction 

studies: (1) insufficient monitoring following reintroduction 

(usually ceasing after 4 years); (2) inadequate documentation, 

which is especially acute for reintroductions that are regarded 

as failures; (3) lack of understanding of the underlying reasons 

for decline in existing plant populations; (4) overly optimistic evaluation of success based on short-term results; and (5) 

poorly defined success criteria for reintroduction projects. We 

therefore conclude that the value of plant reintroductions as a 

conservation tool could be improved by: (1) an increased focus 

on species biology; (2) using a higher number of transplants 

(preferring seedlings rather than seeds); (3) taking better account of seed production and recruitment when assessing the 

success of reintroductions; (4) a consistent long-term monitoring after reintroduction.

Link: http://www.researchgate.net/publication/229104055_

How_successful_are_plant_species_reintroductions/file/

d912f513efbd06d991.pdf

414. Gray LK. 2011. Assisted migration to address 

climate change: recommendations for reforestation in western Canada [Dissertation]. Edmonton, Alberta, Canada: 

University of Alberta. 229 p.

Type: Dissertation

Geographic Area: Western Canada

Compilers’ Keywords: tree populations, seed sources, forest 

management

Abstract: A changing climate is the largest threat to forest 

productivity in western Canada and to the ability of forested 

landscapes to provide ecological and economic services, both 

now and in the future. As climate changes, locally adapted tree 

populations become mismatched with local conditions, leading to mal-adaptation that may result in a reduction in forest 

health and productivity. This problem can be reduced with 

interventions that match reforestation stock to anticipated future environments. As such, there is a pressing need to inform 

such actions by carefully developing and contextualizing scientific information and by applying it to provincial reforestation policies. Assisted migration is a climate change adaptation 

strategy used in the forestry sector, where species and seed 

412. Freeman DC, Turner WA, McArthur ED, 

Graham JH. 1991. Characterization of a narrow hybrid zone 

between two subspecies of big sagebrush. American Journal of 

Botany 78:805-815.

Type: Journal 

Geographic Area: Utah, USA

Compilers’ Keywords: Artemisia tridentata, morphological 

variation

Abstract: Hybridization between Artemisia tridentata ssp. 

tridentata and A. t. ssp. vaseyana occurs in a narrow elevational zone along the west face and canyons of the Wasatch 

Mountains of Utah. Two sites in central Utah (Orem and Salt 

Creek) were examined. The parental taxa differ in anatomy, 

morphology, flavonoid and coumarin content, as well as various growth parameters. Hybrids are intermediate for the majority of traits examined, including flavonoid and coumarin 

composition. Multivariate statistics show that these taxa can 

be unambiguously identified using simple field measures of 

morphology. A hybrid index was constructed using principal 

component analyses on morphological variables.

Link: http://facultyweb.berry.edu/jgraham/Wang%20Narrow%204%20Rec.pdf

413. Godefroid S, Piazza C, Rossi G, Buord S, 

Stevens A, Aguraiuja R, Cowell C, Weekley CW, 

Vogg G, Iriondo JM, Johnson I, Dixon B, Gordon 

DR, Magnanon S, Valentin B, Bjureke K, Koopman 

R, Vicens M, Virevaire M, Vanderborght T. 

2011. How successful are plant species reintroductions?

Biological Conservation 144:672-682.

Type: Journal 

Geographic Area: Global

Keywords: species translocation, population reinforcement, 

population supplementation, population augmentation, restored populations

Abstract: Reintroduction of native species has become increasingly important in conservation worldwide for recovery 

of rare species and restoration purposes. However, few studies have reported the outcome of reintroduction efforts in 

plant species. Using data from the literature combined with 

a questionnaire survey, this paper analyses 249 plant species 

reintroductions worldwide by assessing the methods used and 

the results obtained from these reintroduction experiments. 

The objectives were: (1) to examine how successful plant 

species reintroductions have been so far in establishing or significantly augmenting viable, self-sustaining populations in 

nature; (2) to determine the conditions under which we might 

expect plant species reintroductions to be most successful; (3) 

to make the results of this survey available for future plant 

reintroduction trials. Results indicate that survival, flowering 

and fruiting rates of reintroduced plants are generally quite low 

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north shift is equivalent to approximately 44 m upward shift 

in elevation. However, additional non-climatic factors should 

be considered when matching reforestation stock to suitable 

planting environments. 

Link: http://www.ualberta.ca/~ahamann/publications/pdfs/

Gray_Hamann_2012.pdf

416. Gray LK, Gylander T, Mbogga MS, Chen P, 

Hamann A. 2011. Assisted migration to address climate 

change: recommendations for aspen reforestation in western 

Canada. Ecological Applications 21:1591-1603.

Type: Journal 

Geographic Area: Western Canada

Keywords: bioclimate envelope modeling, climate change, 

ecological genetics, reforestation, remote sensing, seed transfer guidelines, seed zones

Abstract: Human-aided movement of species populations in 

large-scale reforestation programs could be a potent and costeffective climate change adaptation strategy. Such large-scale 

management interventions, however, tend to entail the risks of 

unintended consequences, and we propose that three conditions should be met before implementing assisted migration 

in reforestation programs: (1) evidence of a climate related 

adaptation lag, (2) observed biological impacts, and (3) robust 

model projections to target assisted migration efforts. In a case 

study of aspen (Populus tremuloides Michaux.) we use reciprocal transplant experiments to study adaptation of tree populations to local environments. Second, we monitor natural aspen 

populations using the MODIS enhanced vegetation index as 

a proxy for forest health and productivity. Last, we report results from bioclimate envelope models that predict suitable 

habitat for locally adapted genotypes under observed and predicted climate change. The combined results support assisted 

migration prescriptions and indicate that the risk of inaction 

likely exceeds the risk associated with changing established 

management practices. However, uncertainty in model projections also implies that we are restricted to a relatively short 

20-year planning horizon for prescribing seed movement in 

reforestation programs. We believe that this study exemplifies 

a safe and realistic climate change adaptation strategy based on 

multiple sources of information and some understanding of the 

uncertainty associated with recommendations for assisted migration. Ad hoc migration prescriptions without a similar level 

of supporting information should be avoided in reforestation 

programs.

Link: http://ualberta.ca/~ahamann/publications/pdfs/Gray_

et_al_2011.pdf

417. Green BS, Gardner C, Linnane A, Hawthorne 

PJ. 2010. The good, the bad and the recovery in an assisted 

migration. PLoS ONE 5:e14160.

Type: Journal 

sources are moved to new locations. The goal of this thesis 

is to develop a methodological framework to guide assisted 

migration efforts for forest trees in western Canada, under a 

comprehensive range of future climate projections. To assist 

with these management needs I create a new ecosystem-based 

climate envelope modeling approach for 16 commercially important tree species. Habitat projections show populations already geographically lag behind their optimal climate and the 

magnitude of this lag is projected to double for the 2020s. The 

most pronounced habitat shifts are projected to occur in the boreal forests and the Rocky Mountains, predominately affecting 

black spruce, tamarack, white spruce and aspen populations. 

In a case study for Alberta, I find that genotypes of species 

that are adapted to drier climatic conditions will be the preferred planting stock over much of the commercially managed 

boreal forest. Interestingly, no alternate non-native species to 

Alberta that were examined in this study can be recommended 

with any confidence as planting stock. Finally, I observe high 

uncertainty in projections of suitable habitat for most species 

making reforestation planning beyond the 2050s difficult. 

Using genetic and remote sensing data for aspen populations, 

I show that habitat projections from climate envelope models 

under observed climate change conform well to empirical data 

on loss of aspen productivity and genetic data on sub-optimal 

growth due to mal-adaptation.

415. Gray LK, Hamann A. 2013. Tracking suitable habitat for tree populations under climate change in western North 

America. Climatic Change 117:289-303.

Type: Journal 

Geographic Area: Western North America

Compilers’ Keywords: transfer guidelines, assisted migration, bioclimate envelope modeling, Random Forest, optimal 

climate

Abstract: An important criticism of bioclimate envelope 

models is that many wide-ranging species consist of locally 

adapted populations that may all lag behind their optimal climate habitat under climate change, and thus should be modeled 

separately. Here, we apply a bioclimate envelope model that 

tracks habitat of individual populations to estimate adaptation 

lags for 15 wide-ranging forest tree species in western North 

America. An ensemble classifier modeling approach (Random 

Forest) was used to spatially project the climate space of tree 

populations under observed climate trends (1970s to 2000s) 

and multi-model projections for the 2020s, 2050s, and 2080s. 

We find that, on average, populations already lag behind their 

optimal climate niche by approximately 130 km in latitude, or 

60 m in elevation. For the 2020s we expect an average lag of 

approximately 310 km in latitude or 140 m in elevation, with 

the most pronounced geographic lags in the Rocky Mountains 

and the boreal forest. We show that our results could in principle be applied to guide assisted migration of planting stock in 

reforestation programs using a general formula where 100 km 

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 149

were translocations of threatened, endangered, or sensitive 

species (46 percent). Knowledge of habitat quality, location 

of release area within the species range, number of animals 

released, program length, and reproductive traits allowed 

correct classification of 81 percent of observed translocations as successful or not. 

Link: http://lynx.uio.no/lynx/ibelynxco/04_library/4_3_

publications/G/Griffith_et_al_1989_Translocation_as_a_species_conservation_tool.pdf

419. Guerrant EO Jr. 2012. Characterizing two decades of 

rare plant reintroductions. Chapter 2. In: Maschinski J, Haskins 

KE, editors. Plant Reintroduction in a Changing Climate: 

Promises and Perils. Washington (DC): Island Press. p 9-29. 

Type: Book Section

Geographic Area: Global

Compilers’ Keywords: migration, endangered species, rare 

plant conservation, review

Summary: This chapter presents an overview of reintroduction projects that were documented in the CPC International 

Reintroduction Registry. The goal of the registry and database 

is to assist the conservation community in its ongoing efforts 

to advance the science and improve the practice of conservation. The effort is ongoing, and readers are invited to provide 

information about additional projects by accessing the registry 

on the Center for Plant Conservation website. Any reported 

projects can provide additional insights to the practice of reintroduction, its successes and failures. 

420. Guerrant EO Jr, Kaye TN. 2007. Reintroduction 

of rare and endangered plants: common factors, questions and 

approaches. Australian Journal of Botany 55:362-370.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: review, species extinction, 

conservation

Abstract: The science of reintroduction for conservation 

purposes is young, and there is still much to learn about the 

practice. As a means to achieving biological goals of successfully establishing new populations to enhance a species survival prospects, and project goals, such as learning how to go 

about establishing new populations, reintroduction projects are 

best done as well designed scientific experiments that test explicit hypotheses. Focusing on a range of factors common to 

any reintroduction, we review several empirical reintroduction 

projects with respect to hypotheses tested, experimental materials and methods employed, and evaluate their success in both 

biological and project terms.

Link: http://appliedeco.org/reports/Guerrant%20and%20

Kaye%202007.pdf

Geographic Area: Australia

Compilers’ Keywords: animal migration, biomass, conservation, ecosystem, fisheries, geography, Palinuridae, rock lobster, Jasus edwardsii, population dynamics, Tasmania

Abstract: Assisted migration or translocation of species to 

ameliorate effects of habitat loss or changing environment 

is currently under scrutiny as a conservation tool. A large 

scale experiment of assisted migration over hundreds of kilometres was tested on a morph from a commercial fishery 

of southern rock lobster Jasus edwardsii, to enhance depleted 

populations, improve the yield and sustainability of the fishery, and test resilience to a changing climate. Approximately 

10,000 lower-valued, pale-coloured lobsters were moved 

from deep water to inshore sites (2 in Tasmania [TAS] and 

2 in South Australia [SA]) where the high-value, red morph 

occurs. In TAS this was a northwards movement of 1 degree 

latitude. Growth was measured only in TAS lobsters, and reproductive status was recorded in lobsters from all locations. 

Pale females (TAS) grew 4 times faster than resident pale 

lobsters from the original site and twice as fast as red lobsters 

at their new location. Approximately 30% of translocated 

pale lobsters deferred reproduction for one year after release 

(SA and TAS), and grew around 1 mm yr-1 less compared to 

translocated pale lobsters that did not defer reproduction. In 

spite of this stress response to translocation, females that deferred reproduction still grew 2-6 mm yr-1 more than lobsters 

at the source site. Lobsters have isometric growth whereby 

volume increases as a cube of length. Consequently despite 

the one-year hiatus in reproduction, increased growth increases fecundity of translocated lobsters, as the increase in 

size provided a larger volume for producing and incubating 

eggs in future years. Assisted migration improved egg production and growth, despite a temporary stress response, and 

offers a tool to improve the production, sustainability and 

resilience of the fishery.

Link: http://www.ncbi.nlm.nih.gov/pubmed/21151965

418. Griffith B, Scott JM, Carpenter JW, Reed C. 

1989. Translocation as a species conservation tool: status and 

strategy. Science 245:477-780.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: review, migration, wildlife, endangered species, surveys

Abstract: Surveys of recent (1973 to 1986) intentional releases of native birds and mammals to the wild in Australia, 

Canada, Hawaii, New Zealand, and the United States were 

conducted to document current activities, identify factors 

associated with success, and suggest guidelines for enhancing future work. Nearly 700 translocations were conducted 

each year. Native game species constituted 90 percent of 

translocations and were more successful (86 percent) than 

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organisms to climate change. In essence, one divides the rate 

of climate change by the rate of spatial climate variability to 

obtain a speed at which species must migrate over the surface 

of the earth to maintain constant climate conditions. However, 

to apply the algorithm for conservation and management purposes, additional information is needed to improve realism at 

local scales. For example, destination information is needed to 

ensure that vectors describing speed and direction of required 

migration do not point toward a climatic cul-de-sac by pointing beyond mountain tops. Here, we present an analytical approach that conforms to standard velocity algorithms if climate 

equivalents are nearby. Otherwise, the algorithm extends the 

search for climate refugia, which can be expanded to search 

for multivariate climate matches. With source and destination 

information available, forward and backward velocities can be 

calculated allowing useful inferences about conservation of 

species (present-to-future velocities) and management of species populations (future-to-present velocities).

Link: http://www.ncbi.nlm.nih.gov/pubmed/25310933

423. Hampe A. 2011. Plants on the move: the role of seed 

dispersal and initial population establishment for climate-driven 

range expansions. Acta Oecologica 37:666-673.

Type: Journal 

Geographic Area: Global

Keywords: biotic interactions, dispersal pathways, leading edge, long-distance seed dispersal, range dynamics, tree 

recruitment

Abstract: Recent climate change will presumably allow 

many plant species to expand their geographical range up to 

several hundred kilometres towards the poles within a few 

decades. Much uncertainty exists however to which extent 

species will actually be able to keep pace with a rapidly changing climate. A suite of direct and indirect research approaches 

have explored the phenomenon of range expansions, and the 

existing evidence is scattered across the literature of diverse 

research subdisciplines. Here I attempt to synthesise the available information within a population ecological framework in 

order to evaluate implications of patterns of seed dispersal and 

initial population establishment for range expansions. After introducing different study approaches and their respective contributions, I review the empirical evidence for the role of longdistance seed dispersal in past and ongoing expansions. Then 

I examine how some major ecological determinants of seed 

dispersal and colonisation processes—population fecundity, 

dispersal pathways, arrival site conditions, and biotic interactions during recruitment—could be altered by a rapidly changing climate. While there is broad consensus that long-distance 

dispersal is likely to be critical for rapid range expansions, it 

remains challenging to relate dispersal processes and pathways 

with the establishment of pioneer populations ahead of the 

continuous species range. Further transdisciplinary efforts are 

421. Hamann A, Aitken SN. 2012. Conservation planning under climate change: accounting for adaptive potential 

and migration capacity in species distribution models. Diversity 

and Distributions 19:268-280.

Type: Journal 

Geographic Area: Western Canada

Keywords: bioclimate envelope models, climate change, conservation, genetic adaptation, migration, niche models

Abstract: A number of assumptions underpinning the use of 

species distribution models to predict biological responses to 

climate change are violated for temperate and boreal tree species that are widespread, long-lived and genetically adapted to 

local climate conditions. To address this situation, we propose 

a methodology to account for the potential effects of genetic 

structure, adaptive potential and limited migration capacity. 

Similar to the widely used ‘no migration’ and ‘unlimited migration’ scenarios, we employ more refined biological response 

scenarios to evaluate the potential effects of genetic adaptation 

to local environments and the capacity of species to adapt and 

migrate. These scenarios are realized by two sets of geographic 

delineations that partition the species range into multiple populations and that subdivide the study area into smaller landscape 

units. In a case study for British Columbia, we demonstrate 

how the approach can be used to evaluate the adequacy of a 

reserve system of 90 protected areas to ensure long-term maintenance of forest genetic resources for 48 tree species. We find 

that between 35% and 85% of locally adapted populations in 

protected areas are maintained under a median climate change 

scenario until the end of the century. A sensitivity analysis 

shows that assumptions about migration and adaptation capacity of species have a major effect on the projected conservation 

status. We propose that the results of species distribution models have practical value for conservation planning if the focus 

on maintenance rather than loss of suitable habitat. Accounting 

for genetic structure, adaptive potential and migration capacity 

through best-case and worst-case scenarios provide important 

information to effectively allocate limited resources available 

for conservation action. 

Link: http://www.ualberta.ca/~ahamann/publications/pdfs/

Hamann_Aitken_2013.pdf

422. Hamann A, Roberts DR, Barber QE, Carroll 

C, Nielsen SE. 2014. Velocity of climate change algorithms 

for guiding conservation and management. Global Change 

Biology 21:997-1004.

Type: Journal 

Geographic Area: North America

Keywords: assisted migration, climate change adaptation, climate change vulnerability, conservation, no-analogue climates

Abstract: The velocity of climate change is an elegant analytical concept that can be used to evaluate the exposure of 

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dispersal estimates of sympatric populations of 10 tropical tree 

species. We re-analysed 26 data sets consisting of mapped allozyme, SSR (simple sequence repeat), RAPD (random amplified polymorphic DNA) or AFLP (amplified fragment length 

polymorphism) genotypes from two rainforest sites in French 

Guiana. Gene dispersal estimates were obtained for at least 

one marker in each species, although the estimation procedure 

failed under insufficient marker polymorphism, limited sample 

size, or inappropriate sampling area. Estimates generally suffered low precision and were affected by assumptions regarding the effective population density. Averaging estimates over 

data sets, the extent of gene dispersal ranged from 150 m to 

1200 m according to species. Smaller gene dispersal estimates 

were obtained in species with heavy diaspores, which are presumably not well dispersed, and in populations with high local adult density. We suggest that limited seed dispersal could 

indirectly limit effective pollen dispersal by creating higher 

local tree densities, thereby increasing the positive correlation 

between pollen and seed dispersal distances. We discuss the 

potential and limitations of our indirect estimation procedure 

and suggest guidelines for future studies.

Link: http://www.ncbi.nlm.nih.gov/pubmed/16448421

426. Hereford J. 2009. A quantitative survey of local adaptation and fitness trade-offs. American Naturalist 173:579-588.

Type: Journal 

Geographic Area: Global

Keywords: genetic drift, genotype-by-environment interaction, local adaptation, natural selection, population divergence, 

reciprocal transplant

Abstract: The long history of reciprocal transplant studies 

testing the hypothesis of local adaptation has shown that populations are often adapted to their local environments. Yet many 

studies have not demonstrated local adaptation, suggesting that 

sometimes native populations are no better adapted than are 

genotypes from foreign environments. Local adaptation may 

also lead to trade-offs, in which adaptation to one environment comes at a cost of adaptation to another environment. 

I conducted a survey of published studies of local adaptation 

to quantify its frequency and magnitude and the costs associated with local adaptation. I also quantified the relationship 

between local adaptation and environmental differences and 

the relationship between local adaptation and phenotypic divergence. The overall frequency of local adaptation was 0.71, 

and the magnitude of the native population advantage in relative fitness was 45%. Divergence between home site environments was positively associated with the magnitude of local 

adaptation, but phenotypic divergence was not. I found a small 

negative correlation between a population’s relative fitness in 

its native environment and its fitness in a foreign environment, 

indicating weak trade-offs associated with local adaptation. 

These results suggest that populations are often locally adapted 

clearly needed to address this link, key for understanding how 

plant populations ’move’ across changing landscapes.

Link: http://www.sciencedirect.com/science/article/pii/

S1146609X11000749

424. Hampe A, Petit RJ. 2005. Conserving biodiversity 

under climate change: the rear edge matters. Ecology Letters 

8:461-467.

Type: Journal 

Geographic Area: Global

Keywords: demography, diversification, extinction, genetic 

differentiation, glacial refugia, global change, leading edge, 

peripheral populations, Quaternary, trailing edge

Abstract: Modern climate change is producing poleward 

range shifts of numerous taxa, communities and ecosystems 

worldwide. The response of species to changing environments 

is likely to be determined largely by population responses at 

range margins. In contrast to the expanding edge, the low-latitude limit (rear edge) of species ranges remains understudied, 

and the critical importance of rear edge populations as longterm stores of species genetic diversity and foci of speciation 

has been little acknowledged. We review recent findings from 

the fossil record, phylogeography and ecology to illustrate that 

rear edge populations are often disproportionately important for 

the survival and evolution of biota. Their ecological features, 

dynamics and conservation requirements differ from those of 

populations in other parts of the range, and some commonly 

recommended conservation practices might therefore be of 

little use or even counterproductive for rear edge populations.

Link: http://eeslmu.de/eeswiki/images/RearEdgeEcolLetters05.pdf

425. Hardy OJ, Maggia L, Bandou E, Breyne P, 

Caron H, Chevallier MH, Doligez A, Dutech C, 

Kremer A, Latouche-Halle C, Troispoux V, Veron 

V, Degen B. 2006. Fine-scale genetic structure and gene 

dispersal inferences in 10 neotropical tree species. Molecular 

Ecology 15:559-571.

Type: Journal 

Geographic Area: South America

Keywords: French Guiana, gene dispersal, seed dispersal, 

spatial genetic structure, tropical trees

Abstract: The extent of gene dispersal is a fundamental factor 

of the population and evolutionary dynamics of tropical tree 

species, but directly monitoring seed and pollen movement is 

a difficult task. However, indirect estimates of historical gene 

dispersal can be obtained from the fine-scale spatial genetic 

structure of populations at drift-dispersal equilibrium. Using 

an approach that is based on the slope of the regression of 

pairwise kinship coefficients on spatial distance and estimates 

of the effective population density, we compare indirect gene 

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152 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

by such long-lived sessile organisms. The rate of climate 

change forecast for the future is 10–100 times faster than the 

rate of deglacial warming. Unless steps are taken to facilitate 

the migratory response of organisms to the forecast changes, 

then widespread extinction is likely. Artificial dispersal of 

trees and other organisms of limited dispersal and/or migratory capacity, the general extension of the legal protection 

currently afforded to some threatened organisms only within 

designated reserves, and the integration of wildlife habitat 

requirements and of wildlife corridors into human landscape 

utilization are all likely to be necessary. Stringent measures 

to limit the extent of future climate change by limiting emissions of greenhouse gases will also be necessary if the possibility of widespread and even catastrophic extinction is to 

be avoided.

Link: http://download.bioon.com.cn/view/upload/month_10

06/20100628_92d6aa5f6c76c0ba6913YYtpqqnBmHzn.attach.

pdf

429. Huntley B. 1991. Historical lessons for the future. 

Chapter 19. In: Spellerberg IF, Goldsmith FB, Morris MK, 

editors. Scientific Management of Temperate Communities for 

Conservation. Blackwell, Oxford: British Ecological Society. 

p 473-503. 

Type: Book Section

Geographic Area: Global

Compilers’ Keywords: climate change, conservation, succession, pollen, vegetation, species distribution, migration, 

review

Introduction (end paragraph): Chapter presents examples illustrating the use of palynological data at various spatial scales, and with varying temporal resolution, 

to address conservation problems. They show evidence of 

changes in communities at various scales, of the degree of 

permanence of particular communities, of the former character of the vegetation in particular localities, of the transient 

nature of some past communities and of the mechanisms of 

response to broad-scale environmental changes. Because 

few examples of small-scale palynological studies that address conservation problems have been published, and because many conservation efforts will continue to be directed 

towards individuals sites, account are given of several such 

studies. In contrast, many of the broad-scale results have already been published and so only the major conclusions are 

presented here. However, it is from these conclusions that 

the most important implications for conservation arise, calling into question current strategies that focus primarily upon 

the establishment of nature reserves at sites that are today 

occupied by particular organisms or communities. It is also 

at the broad scale that palynological data can contribute to 

current research related to global environmental change and 

to global conservation. 

but stochastic processes such as genetic drift may limit the efficacy of divergent selection.

Link: http://www.webpages.uidaho.edu/~snuismer/Nuismer_

Lab/548_readings_files/Hereford%202009.pdf

427. Hu FS, Hampe A, Petit RJ. 2009. Paleoecology 

meets genetics: deciphering past vegetational dynamics.

Frontiers in Ecology and the Environment 7:371-379.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: paleoecology, migration, Quaternary, 

climate change, tree distribution, refugia, genetic conservation

Abstract: Genetic analysis adds a novel dimension to paleoecology that is becoming increasingly important for elucidating vegetational dynamics in relation to climate change. 

Because past vegetational changes have often left distinct 

genetic imprints on current plant populations, genetic analysis can add detail to fossil-based reconstructions. Recent 

surveys of DNA polymorphisms yield new insights into past 

vegetational changes and address long-standing paleoecological questions. These studies reveal that small tree populations survived in mid- to high-latitude refugia throughout 

the Quaternary glacial episodes. They further reveal migration pathways and demographic processes during postglacial 

range expansion. These results challenge previous notions 

regarding tree species responses to climate change and help 

to identify genetic conservation targets. Here, we review 

these recent advances and outline research prospects at the 

interface between paleoecology and genetics.

Link: http://www.esajournals.org/doi/abs/10.1890/070160

428. Huntley B. 1991. How plants respond to climate 

change: migration rates, individualism and the consequences for 

plant communities. Annals of Botany 67:15-22.

Type: Journal 

Geographic Area: Global

Keywords: climate change, pollen analysis, Quaternary, migration, individualism, plant communities

Abstract: The magnitude of climate changes forecast for 

the next century is comparable to the magnitude of warming 

during the last deglaciation. No climate change of similar 

magnitude has occurred since that event. The palaeoecological evidence of the response, especially of plants, to 

past climate change indicates that evolutionary adaptation 

has played no more than a minor role and that migration is 

the usual response of organisms to climate change. The individualism of response has important implications with respect to changes in the nature of vegetation and ecosystems. 

The maximum realized rates of migratory response by trees, 

although perhaps matching the maximum potential rates, are 

close to the maximum that it is believed can be achieved 

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CHAPTER 3 • MIGRATION • RESEARCH

Abstract: We evaluate genetic test plantations of North 

American Douglas-fir provenances in Europe to quantify how 

tree populations respond when subjected to climate regime 

shifts, and we examined whether bioclimate envelope models 

developed for North America to guide assisted migration under 

climate change can retrospectively predict the success of these 

provenance transfers to Europe. The meta-analysis is based 

on long-term growth data of 2,800 provenances transferred to 

120 European test sites. The model was generally well suited 

to predict the best performing provenances along north-south 

gradients in Western Europe, but failed to predict superior performance of coastal North American populations under continental climate conditions in Eastern Europe. However, model 

projections appear appropriate when considering additional 

information regarding adaptation of Douglas-fir provenances 

to withstand frost and drought, even though the model partially 

fails in a validation against growth traits alone. We conclude by 

applying the partially validated model to climate change scenarios for Europe, demonstrating that climate trends observed 

over the last three decades warrant changes to current use of 

Douglas-fir provenances in plantation forestry throughout 

Western and Central Europe.

Link: http://www.ncbi.nlm.nih.gov/pubmed/24737595

432. Iverson LR, Prasad AM. 1998. Predicting abundance of 80 tree species following climate change in the eastern 

United States. Ecological Monographs 68:465-485.

Type: Journal 

Geographic Area: Eastern USA

Keywords: climate change, envelope analysis, forest inventory, geographic information systems (GIS), global change, 

landscape ecology, predictive vegetation mapping, regression 

tree analysis (RTA), species—environment relationships, tree 

species distribution, tree species migration, tree species ranges

Abstract: Projected climate warming will potentially have 

profound effects on the earth’s biota, including a large redistribution of tree species. We developed models to evaluate 

potential shifts for 80 individual tree species in the eastern 

United States. First, environmental factors associated with 

current ranges of tree species were assessed using geographic 

information systems (GIS) in conjunction with regression 

tree analysis (RTA). The method was then extended to better 

understand the potential of species to survive and/or migrate 

under a changed climate. We collected, summarized, and analyzed data for climate, soils, and use, elevation, and species 

assemblages for > 2,100 counties east of the 100th meridian. 

Forest Inventory Analysis (FIA) data for > 100,000 forested 

plots in the East provided the tree species range and abundance 

information for the trees. RTA was used to devise prediction 

rules from current species–environment relationships, which 

were then used to replicate the current distribution as well as 

predict the future potential distributions under two scenarios 

430. Ibanez I, Clark JS, Dietze MC. 2008. Evaluating 

the sources of potential migrant species: implications under climate change. Ecological Applications 18:1664-1678.

Type: Journal 

Geographic Area: Southeastern USA

Keywords: climate change, climate envelope, migration, 

seedling recruitment, source of migrant species, survival, 

transplant experiment, tree species

Abstract: As changes in climate become more apparent, 

ecologists face the challenge of predicting species responses 

to the new conditions. Most forecasts are based on climate 

envelopes (CE), correlative approaches that project future distributions on the basis of the current climate often assuming 

some dispersal lag. One major caveat with this approach is 

that it ignores the complexity of factors other than climate that 

contribute to a species’ distributional range. To overcome this 

limitation and to complement predictions based on CE modeling we carried out a transplant experiment of resident and 

potential-migrant species. Tree seedlings of 18 species were 

planted side by side from 2001 to 2004 at several locations in 

the southern Appalachians and in the North Carolina Piedmont 

(USA). Growing seedlings under a large array of environmental conditions, including those forecasted for the next decades, 

allowed us to model seedling survival as a function of variables characteristic of each site, and from here we were able 

to make predictions on future seedling recruitment. In general, 

almost all species showed decreased survival in plots and years 

with lower soil moisture, including both residents and potential migrants, and in both locations, the southern Appalachians 

and the Piedmont. The detrimental effects that anticipated arid 

conditions could have on seedling recruitment contradict some 

of the projections made by CE modeling, where many of the 

species tested are expected to increase in abundance or to expand their ranges. These results point out the importance of 

evaluating the potential sources of migrant species when modeling vegetation response to climate change, and considering 

that species adapted to the new climate and the local conditions 

may not be available in the surrounding regions.

Link: http://www.srs.fs.usda.gov/pubs/ja/2008/ja_2008_

ibanez_001.pdf

431. Isaac-Renton MG, Roberts DR, Hamann A, 

Spiecker H. 2014. Douglas-fir plantations in Europe: a retrospective test of assisted migration to address climate change. 

Global Change Biology 20:2607-2617.

Type: Journal 

Geographic Area: Europe

Keywords: bioclimatic envelope models, exotic species, 

model validation, no-analogue climates, provenance trials, 

Pseudotsuga menziesii, species distribution models
154 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

CHAPTER 3 • MIGRATION • RESEARCH

low and high colonization probabilities as a result of low 

and high densities of target trees (high source strength) as 

well as high densities of forest (high sink strength). We also 

assess migration potential for species by concentric rings 

around the current boundary. Model outputs show the generally limited nature of migration for all five species over 100 

years. There is a relatively high probability of colonization 

within a zone of 10–20 km (depending on habitat quality and 

species abundance) from the current boundary, but a small 

probability of colonization where the distance from the current boundary exceeds about 20 km. Whether biologically 

plausible or not, rare very long distance migration events are 

not sufficient to rescue migration. Species abundance (the 

source strength of migration) near the range boundary carried relatively more influence than percentage forest cover 

(sink strength) in determining migration rates. The transect 

evaluation revealed the importance of abundance of the species near the boundary, indicating that rare species may have 

much more difficulty in unassisted northward migration 

due to climate change. The concentric rings analysis of the 

model outputs showed that only the first 10–20 km of area 

would have a reasonably high probability of colonization. 

Rare, long-distance events permit colonization of remote 

outliers, but much more needs to be understood about the 

likelihood of these rare events to predict the frequency of 

outlier establishment.

Link: http://www.des.ucdavis.edu/faculty/mschwartz/Website%20publications/IversonGlobalEcol2004.pdf

434. Iverson LR, Prasad AM, Matthews SN, 

Peters MP. 2011. Lessons learned while integrating habitat, 

dispersal, disturbance, and life-history traits into species habitat 

models under climate change. Ecosystems 14:1005-1020.

Type: Journal 

Geographic Area: Eastern USA

Keywords: climate change, eastern United States, randomForest, statistical modeling, migration, trees, birds, DISTRIB, 

SHIFT, ModFacs 

Abstract: We present an approach to modeling potential 

climate-driven changes in habitat for tree and bird species in 

the eastern United States. First, we took an empirical-statistical modeling approach, using RandomForest, with species 

abundance data from national inventories combined with soil, 

climate, and landscape variables, to build abundance-based 

habitat models for 134 tree and 147 bird species. We produced 

lists of species for which suitable habitat tends to increase, 

decrease, or stay the same for any region. Independent assessments of trends of large trees versus seedlings across the 

eastern U.S. show that 37 of 40 species in common under 

both studies are currently trending as modeled. We developed 

framework, ModFacs, in which we used the literature to assign 

default modification factor scores for species characteristics 

of climate change with twofold increases in the level of atmospheric CO2. Validation measures prove the utility of the 

RTA modeling approach for mapping current tree importance 

values across large areas, leading to increased confidence in 

the predictions of potential future species distributions. With 

our analysis of potential effects, we show that roughly 30 species could expand their range and/or weighted importance at 

least 10%, while an additional 30 species could decrease by 

at least 10%, following equilibrium after a changed climate. 

Depending on the global change scenario used, 4–9 species 

would potentially move out of the United States to the north. 

Nearly half of the species assessed (36 out of 80) showed the 

potential for the ecological optima to shift at least 100 km to 

the north, including seven that could move > 250 km. Given 

these potential future distributions, actual species redistributions will be controlled by migration rates possible through 

fragmented landscapes.

Link: http://www.clas.ufl.edu/users/mbinford/GEOXXXX_

Biogeography/LiteratureForLinks/iverson_and_prasad_1998_

predicting_abundance_80_tree_species_climate_change_

EUS_EcolMonogr.pdf

433. Iverson LR, Schwartz MW, Prasad AM. 2004. 

How fast and far might tree species migrate in the eastern United 

States due to climate change? Global Ecology and Biogeography 

13:209-219.

Type: Journal 

Geographic Area: Eastern USA

Keywords: climate change, Diospyros virginiana, fragmented 

habitat, global warming, Liquidambar styraciflua, migration, 

Oxydendrum arboretum, Pinus taeda, Quercus falcate, United 

States

Abstract: We describe and use a model, SHIFT, to estimate 

potential migration due to climate change over the next 100 

years in the eastern United States. Five species, currently 

confined to the eastern half of the United States and not extending into Canada, were used to assess migration potential: 

Diospyros virginiana (persimmon), Liquidambar styraciflua

(sweetgum), Oxydendrum arboreum (sourwood), Pinus taeda (loblolly pine), and Quercus falcata var. falcata (southern 

red oak). SHIFT is a matrix simulation model using simple 

inverse power functions to provide a distance decay of seed 

dispersal and is driven primarily by the abundance of the species near the boundary, the forest density within and beyond 

the boundary, and the distance between cells. For each cell 

outside the current boundary, the model creates an estimate 

of the probability that each unoccupied cell will become 

colonized over a period of 100 years. SHIFT is a ‘fat-tailed’ 

migration model that allows rare very long distance dispersal events and colonization could occur up to 500 km beyond the current distribution boundary. Model outputs were 

analysed using transects through sections showing relatively 
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 155

436. Jump AS, Matyas C, Penuelas J. 2009. The 

altitude-for-latitude disparity in the range retractions of woody 

species. Trends in Ecology and Evolution 24:694-701.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: review, migration, climate change, 

species distribution, ecosystem services, dendroecology, genetic adaptation, provenance trial, phenology, rates, range 

shifts

Abstract: Increasing temperatures are driving rapid upward 

range shifts of species in mountains. An altitudinal range retreat of 10 m is predicted to translate into an approx. 10-km 

latitudinal retreat based on the rate at which temperatures decline with increasing altitude and latitude, yet reports of latitudinal range retractions are sparse. Here, we examine potential 

climatic, biological, anthropogenic and methodological explanations for this disparity. We argue that the lack of reported 

latitudinal range retractions stems more from a lack of research 

effort, compounded by methodological difficulties, rather than 

from their absence. Given the predicted negative impacts of 

increasing temperatures on wide areas of the latitudinal distributions of species, the investigation of range retractions should 

become a priority in biogeographical research.

437. Kaye TN. 2008. Vital steps toward success of endangered plant reintroductions. Native Plants Journal 9:313-322.

Type: Journal 

Geographic Area: Global

Keywords: recovery, rare species, restoration, translocation, 

population, augmentation

Abstract: Reintroduction of endangered plants faces many 

challenges, but the survival of some species may depend on 

its success. What measures should practitioners take to ensure 

a successful project, and how should success be measured? 

Steps in the reintroduction process include planning and 

identification of objectives, finding source material, propagation, site selection, site preparation, outplanting, monitoring, 

evaluation and interpretation, feedback to improve protocols, 

communication with others, habitat maintenance, and repeated 

actions if necessary to meet objectives. Conducting reintroductions as designed experiments and applying the results through 

adaptive management will maximize the effectiveness of 

reintroductions.

Link: http://appliedeco.org/reports/Kaye%202008%20Reintroduction%20Native%20Plants%20Journal.pdf

438. Kelly AE, Goulden ML. 2008. Rapid shifts in distribution with recent climate change. Proceedings of the National 

Academy of Science USA 105:11823-11826.

Type: Journal 

that cannot be readily assessed in such models, including 12 

disturbance factors (for example, drought, fire, insect pests), 

nine biological factors (for example, dispersal, shade tolerance), and assessment scores of novel climates, long-distance 

extrapolations, and output variability by climate model and 

emission scenario. We also used a spatially explicit cellular 

model, SHIFT, to calculate colonization potentials for some 

species, based on their abundance, historic dispersal distances, 

and the fragmented nature of the landscape. By combining 

results from the three efforts, we can create projections of potential climate change impacts over the next 100 years or so. 

Here we emphasize some of the lessons we have learned over 

16 years in hopes that they may help guide future experiments, 

modeling efforts, and management. 

Link: http://www.researchgate.net/publication/225455924_

Lessons_Learned_While_Integrating_Habitat_Dispersal_Disturbance_and_Life-History_Traits_into_Species_Habitat_

Models_Under_Climate_Change/file/9fcfd501955d5a053e.

pdf

435. Jump AS, Penuelas J. 2005. Running to stand still: 

adaptation and the response of plants to rapid climate change. 

Ecology Letters 8:1010-1020.

Type: Journal 

Geographic Area: Global

Keywords: adaptation, climate change, conservation, decreased fitness, extinction risk, gene flow, genetic diversity, 

habitat fragmentation, migration, plant populations

Abstract: Climate is a potent selective force in natural populations, yet the importance of adaptation in the response of 

plant species to past climate change has been questioned. As 

many species are unlikely to migrate fast enough to track the 

rapidly changing climate of the future, adaptation must play an 

increasingly important role in their response. In this paper we 

review recent work that has documented climate-related genetic diversity within populations or on the microgeographical 

scale. We then describe studies that have looked at the potential 

evolutionary responses of plant populations to future climate 

change. We argue that in fragmented landscapes, rapid climate 

change has the potential to overwhelm the capacity for adaptation in many plant populations and dramatically alter their genetic composition. The consequences are likely to include unpredictable changes in the presence and abundance of species 

within communities and a reduction in their ability to resist and 

recover from further environmental perturbations, such as pest 

and disease outbreaks and extreme climatic events. Overall, 

a range-wide increase in extinction risk is likely to result. We 

call for further research into understanding the causes and 

consequences of the maintenance and loss of climate-related 

genetic diversity within populations.

Link: http://www3.botany.ubc.ca/rieseberglab/plantevol/

Jump_and_Penuelas2005.pdf

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156 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

440. Kim M-S, Richardson BA, McDonald GI, 

Klopfenstein NB. 2011. Genetic diversity and structure of 

western white pine (Pinus monticola) in North America: a baseline study for conservation, restoration, and addressing impacts 

of climate change. Tree Genetics & Genomes 7:11-21.

Type: Journal 

Geographic Area: Western USA

Keywords: genetic variation, tree population structure, Pinus 

monticola

Abstract: Western white pine (Pinus monticola) is an economically and ecologically important species in western North 

America that has declined in prominence over the past several 

decades, mainly due to the introduction of Cronartium ribicola 

(cause of white pine blister rust) and reduced opportunities 

for regeneration. Amplified fragment length polymorphism 

(AFLP) markers were used to assess the genetic diversity and 

structure among populations at 15 sites (e.g., provenances) 

across the native range of western white pine. The level of 

genetic diversity was different among 15 populations tested 

using 66 polymorphic AFLP loci. Nei’s gene diversity (HE) 

at the population level ranged from 0.187 to 0.316. Genetic 

differentiation (GST) indicated that 20.1% of detected genetic 

variation was explained by differences among populations. In 

general, populations below 45° N latitude exhibited a higher 

level of genetic diversity than higher latitude populations. 

Genetic distance analysis revealed two major clades between 

northern and southern populations, but other well supported 

relationships are also apparent within each of the two clades. 

The complex relationships among populations are likely derived from multiple factors including migration, adaptation, 

and multiple glacial refugia, especially in higher latitudes. 

Genetic diversity and structure revealed by this study will aid 

recognition and selection of western white pine populations for 

species management and conservation programs, especially in 

consideration of current and future climate changes.

Link: http://www.rmrs.nau.edu/publications/kim_et_

al_2010b/kim_et_al_2010b.pdf

441. King MG, Horning ME, Roalson EH. 2009.

Range persistence during the last glacial maximum: Carex 

macrocephala was not restricted to glacial refugia. Molecular 

Ecology 18:4256-4269.

Type: Journal 

Geographic Area: Northwestern North America

Keywords: coastal refuge, Cyperaceae, Pleistocene glaciation, population genetics

Abstract: The distribution of many species inhabiting northwestern North America has been heavily influenced by the 

climatic changes during the late Pleistocene. Several studies 

have suggested that species were restricted to glacial refugia 

north and/or south of the continental ice sheet front. It is also 

Geographic Area: California, USA

Keywords: plant migration, range shift

Abstract: A change in climate would be expected to shift 

plant distribution as species expand in newly favorable areas 

and decline in increasingly hostile locations. We compared 

surveys of plant cover that were made in 1977 and 2006–2007 

along a 2,314-m elevation gradient in Southern California’s 

Santa Rosa Mountains. Southern California’s climate warmed 

at the surface, the precipitation variability increased, and the 

amount of snow decreased during the 30-year period preceding the second survey. We found that the average elevation of 

the dominant plant species rose by 65 m between the surveys. 

This shift cannot be attributed to changes in air pollution or 

fire frequency and appears to be a consequence of changes in 

regional climate.

Link: http://listmail.energy.ca.gov/sitingcases/genesis_solar/

documents/others/testimony_centr_biological_diversity/

exhibits/Exh.%20816.%20Kelly%20and%20Goulden%20

2008%20%20%20Rapid%20shifts%20in%20plant%20distribution%20with%20recent%20climate%20change.pdf

439. Khasa PD, Dancik BP. 1996. Rapid identification of white-Engelmann spruce by RAPD markers. Theoretical 

Applications of Genetics 92:46-52.

Type: Journal 

Geographic Area: North America

Keywords: hybridization, PCR, Picea, RAPD fingerprints

Abstract: Fragments of random amplified polymorphic 

DNA (RAPDs) were used as markers to distinguish Picea 

glauca (Moench) Voss (white spruce) and Picea engelmannii

Parry (Engelmann spruce). These species and their putative 

hybrids are difficult to differentiate morphologically and 

are collectively known as interior spruce. Four oligodeoxynucleotide decamer primers showed species specific amplification products between white spruce and Engelmann 

spruce. These fragments are highly conserved among seed 

lots and individual trees of each species from diverse geographic origins. The consistency and reproducibility of these 

species-specific amplification products were tested in more 

than two amplification reactions. Therefore, RAPD markers 

can provide genetic markers for easy and rapid identification 

of the specific genetic entry of these spruce species and their 

reported putative hybrids. According to the frequencies of 

the species-specific RAPD markers, it is possible to estimate 

the hybrid fraction, indicative of true introgression between 

the two species. These results are useful for quick identification of both species and their hybrid swarms at any stage 

in the sporophyte phase of the life cycle, for determining 

the occurrence and the magnitude of introgressive hybridization in an overlap zone between the two species, and for 

certification purposes in operational re-forestation and treeimprovement programs.

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 157

productive test sites. Host population effects were limited to 

EMF species abundance rather than species loss, as richness 

per site averaged 15.0 among provenances and did not differ 

by transfer extent (up to 450 km), while Shannon’s diversity 

index declined slightly. Large differences in colonization rates 

of specialist fungi, such as Tomentella stuposa and Clavulina 

cristata, raise the possibility that EMF communities maladapted to soil conditions contributed to the inferior growth of some 

host populations on productive sites. The results of the study 

suggest locally based specificity in host-fungal communities 

is likely a contributing factor in the outcome of provenance 

trials, and should be a consideration in analyzing seed-transfer 

effects and developing strategies for assisted migration. 

443. Kremer A, Ronce O, Robledo-Arnuncio 

JJ, Guillaume F, Bohrer G, Nathan R, Bridle JR, 

Gomulkiewicz R, Klein EK, Ritland K, Kuparinen 

A, Gerber S, Schueler S. 2012. Long-distance gene flow 

and adaptation of forest trees to rapid climate change. Ecology 

Letters 15:378-392.

Type: Journal 

Geographic Area: Global

Keywords: adaptation, climate change, forest trees, gene 

flow, selection

Abstract: Forest trees are the dominant species in many parts 

of the world and predicting how they might respond to climate 

change is a vital global concern. Trees are capable of longdistance gene flow, which can promote adaptive evolution in 

novel environments by increasing genetic variation for fitness. 

It is unclear, however, if this can compensate for maladaptive 

effects of gene flow and for the long-generation times of trees. 

We critically review data on the extent of long-distance gene 

flow and summarize theory that allows us to predict evolutionary responses of trees to climate change. Estimates of longdistance gene flow based both on direct observations and on 

genetic methods provide evidence that genes can move over 

spatial scales larger than habitat shifts predicted under climate 

change within one generation. Both theoretical and empirical 

data suggest that the positive effects of gene flow on adaptation may dominate in many instances. The balance of positive 

to negative consequences of gene flow may, however, differ 

for leading edge, core and rear sections of forest distributions. 

We propose future experimental and theoretical research that 

would better integrate dispersal biology with evolutionary 

quantitative genetics and improve predictions of tree responses 

to climate change.

Link: http://www.ncbi.nlm.nih.gov/pubmed/22372546

444. Kreyling J, Wiesenberg GLB, Thiel D, 

Wohlfart C, Huber G, Walter J, Jentsch A, Konnert 

M, Beierkuhnlein C. 2012. Cold hardiness of Pinus 

nigra Arnold as influenced by geographic origin, warming, and 

hypothesized that the coast of northwestern North America 

could have been a prime location for glacial refugia because 

of the lowering of the eustatic sea level and the concomitant 

rise of the continental shelf because of tectonic rebound. 

Alternatively, some coastal species distributions and demographics may have been unaffected in the long-term by the 

last glacial maximum (LGM). We tested the glacial refugium 

hypothesis on an obligate coastal plant species, Carex macrocephala by sampling 600 individuals from 41 populations with 

11 nuclear microsatellite loci and the rpL16 plastid intragenic 

spacer region. The microsatellite data sets suggest a low level 

of population differentiation with a standardized G’ST = 0.032 

and inbreeding was high with an F = 0.969. The homogenization of the populations along the coast was supported by a principal coordinate analysis, AMOVAS and SAMOVA analyses. 

Analyses using the rpL16 data set support the results of the microsatellite analyses, with a low FST of 0.042. Coalescent and 

mismatch analyses using rpL16 suggest that C. macrocephala 

has not gone through a significant bottleneck within the past 

100, 000 years, although a much earlier population expansion 

was indicated by the mismatch analysis. Carex macrocephala

exhibits the characteristics of metapopulation dynamics and 

on the basis of these results, we concluded that it was not restricted to glacial refugia during the LGM, but that it existed as 

a large metapopulation.

Link: http://www.ncbi.nlm.nih.gov/pubmed/19754517

442. Kranabetter JM, Stoehr MU, O’Neill GA. 

2012. Divergence in ectomycorrhizal communities with foreign Douglas-fir populations and implications for assisted migration. Ecological Applications 22:550-560.

Type: Journal 

Geographic Area: Western Canada

Keywords: climate change mitigation, coevolution, community genetics, foundation species, maladaptation, Southwestern 

British Columbia, Canada

Abstract: Assisted migration of forest trees has been widely 

proposed as a climate change adaptation strategy, but moving tree populations to match anticipated future climates may 

disrupt the geographically based, coevolved association suggested to exist between host trees and ectomycorrhizal fungal 

(EMF) communities. We explored this issue by examining 

the consistency of EMF communities among populations of 

40 year-old Douglas-fir (Pseudotsuga menziesii var. menziesii) 

trees in common-garden field trial using four provenances 

from contrasting coastal climates in southwestern British 

Columbia. Considerable variation in EMF community composition within test sites was found, ranging from 0.38 to 

0.65 in the mean similarity index, and the divergence in EMF 

communities from local populations increased with site productivity. Clinal patterns in colonization success were detected 

for generalist and specialist EMF species on only the two 

CHAPTER 3 • MIGRATION • RESEARCH
158 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Compilers’ Keywords: migration, forest management, assisted migration, Douglas-fir, Pseudotsuga menziesii var. menziesii, loblolly pine, Pinus taeda, conference paper

Abstract: Global climate change and the demand for an alternative, renewable and ecologically friendly source of energy 

significantly increases the role of pine forests in the southeastern United States and population and ecological genomic studies of their adaptive, ecological and economic potential. The 

accelerating global climate change can exceed the adaptive 

potential of pine forests and lead to their contraction on large 

areas. However, some forest tree populations growing now in 

the regions that have ecological conditions similar to the future changes are likely to possess an adaptive potential necessary for spreading adaptation into other regions with similar 

ecological conditions expected in the future. Unfortunately, 

phenotypic plasticity and historically established gene flow 

between remote populations can be insufficient for fast natural 

propagation of genetic adaptations. In this connection the role 

of conservation management and especially that of assisted migration (also called assisted colonization, managed relocation 

or translocation done by physically moving the plant material 

(pollen, seeds and seedlings) to other regions where this material is better adapted to the future environments) increases. 

Modern methods and tools of population, landscape and ecological genomics allow to study the genetic basis of adaptation 

in forest tree species and also detect the main genes responsible for important adaptive and economic traits that can be 

used in assisted gene migration and tree breeding for biomass 

growth, water use efficiency, cellulose content and other traits 

important for adaptation and for bioenergy and biofuel production. The recent data on Douglas-fir (Pseudotsuga menziesii

var. menziesii) and loblolly pine (Pinus taeda L.) populations 

studied for thousands different genes are presented to illustrate 

population and ecological genomic studies in conifers.

Link: https://pag.confex.com/pag/xx/webprogram/Paper1651.

html

446. Krutovsky KV, Burczyk J, Chybicki I, Finkeldey 

R, Pyhäjärvi T, Robledo-Arnuncio JJ. 2012. Gene 

flow, spatial structure, local adaptation, and assisted migration 

in trees. Chapter 4. In: Schnell RJ, Priyadarshan PM, editors. 

Genomics of Tree Crops. New York (NY): Springer New York. 

p 71-116. 

Type: Book Section

Geographic Area: Global

Keywords: assisted migration, gene flow, dispersal, introgression, local adaptation, molecular genetic markers, spatial 

structure

Abstract: Gene flow is a process of transferring and exchanging genetic material among groups of organisms and is an 

important evolutionary factor that greatly affects genetic variation and differentiation in trees. Gene flow within and among 

extreme summer drought. Environmental and Experimental 

Botany 78:99-108.

Type: Journal 

Geographic Area: Europe

Keywords: frost hardiness, black pine, ecotype, cold tolerance, global warming, winter ecology

Abstract: Adaptation to the adverse effects of climate 

change is being investigated more and more through the introduction of species from warmer and drier climates, such 

as the (sub-) Mediterranean Pinus nigra to dry sites in temperate Central Europe. Winter survival, however, may pose 

a serious threat to this strategy as cold extremes, which naturally determine the poleward range limits of forest trees, are 

not expected to follow the general warming trend in the near 

future. Here, juveniles of P. nigra from eight provenances 

throughout Europe were exposed to different climate change 

scenarios (factorial combinations of 42 days of drought and 

warming by 1.6 °C) in a common garden experiment in 

Bayreuth, Germany. Cold hardiness (LT50) was determined 

by the Relative Electrolyte Leakage method (REL) in two 

consecutive winters. Cold hardiness of foliage differed by 

10 °C between the provenances studied and a local adaptation to minimum temperature was found. Cold hardiness 

was further affected by extreme summer drought, increasing cold hardiness by 3.9 °C on average in the subsequent 

winter, and by summer warming, increasing cold hardiness 

by 3.4 °C. Year-round warming had no significant effect on 

cold hardiness. Cold hardiness was related to the content of 

soluble carbohydrates and to the composition of fatty acids 

and alkanes in the needles. Juveniles of P. nigra exhibited 

a comparable cold hardiness as juveniles of species native 

to Central Europe (P. sylvestris, Picea abies, Fagus sylvatica

and Quercus petraea) under the same climatic conditions. 

Cold hardiness of the fine roots of P. nigra averaged -16.5 

°C compared to -23.8 °C on average for needles. Our results 

imply that the cold hardiness of the foliage is adaptive to 

both long-term growing conditions at the seed origin (genetic heritage) and short-term alterations of these conditions 

(individual plasticity), while first hints suggest that cold hardiness of the roots is high and might not be adaptive. For P. 

nigra, below- and above-ground cold hardiness of selected 

provenances in mid-winter appears suitable for cultivation 

in temperate regions.

Link: http://www.sciencedirect.com/science/article/pii/

S0098847211003339

445. Krutovsky KV. 2012. Forest genomics for mitigating climate change and breeding resilient trees. In: International 

Plant and Animal Genome Conference. San Diego (CA): Plant 

and Animal Genome. p 1.

Type: Conference Paper

Geographic Area: Southeastern USA

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 159

to appropriately represented tropical diversity, will allow for 

truly general inference and require the continued use of correlative approaches that may take on increasingly mechanistic 

components. The trade-off between detail and scale is likely to 

characterize the future of global change biodiversity research, 

and birds may be an excellent group to improve, integrate and 

geographically extend current approaches.

Link: http://www.ncbi.nlm.nih.gov/pubmed/20190111

448. Ledig FT, Rehfeldt GE, Jaquish B. 2012.

Projections of suitable habitat under climate change scenarios: 

implications for trans-boundary assisted colonization. American 

Journal of Botany 99:1217-1230.

Type: Journal 

Geographic Area: Western USA

Keywords: assisted colonization, climatic models, conservation, Picea breweriana, Pinaceae, Random Forests

Abstract: Climate change may threaten endemic species with 

extinction, particularly relicts of the Arcto-Tertiary Forest, by 

elimination of their contemporary habitat. Projections of future 

habitat are necessary to plan for conservation of these species. 

We used spline climatic models and modified Random Forests 

statistical procedures to predict suitable habitats for Brewer 

spruce (Picea breweriana), which is endemic to the Klamath 

Region of California and Oregon. We used three general circulation models and two sets of carbon emission scenarios (optimistic and pessimistic) for future climates. Our procedures 

predicted present occurrence of Brewer spruce perfectly. For 

the decades 2030, 2060, and 2090, its projected range within 

the Klamath Region progressively declined, to the point of 

disappearance in the decade 2090. The climate niche was projected to move north to British Columbia, the Yukon Territory, 

and southeastern Alaska. The results emphasize the necessity 

of assisted colonization and trans-boundary movement to prevent extinction of Brewer spruce. The projections provide a 

framework for formulating conservation plans, but planners 

must also consider regulations regarding international plant 

transfers.

Link: http://www.ncbi.nlm.nih.gov/pubmed/22763352

449. Lenoir J, Gegout JC, Marquet PA, de Ruffray 

P, Brisse H. 2008. A significant upward shift in plant species optimum elevation during the 20th century. Science 

320:1768-1771.

Type: Journal 

Geographic Area: Western Europe

Compilers’ Keywords: range shifts, forest plant species, climate change

Abstract: Spatial fingerprints of climate change on biotic 

communities are usually associated with changes in the distribution of species at their latitudinal or altitudinal extremes. 

populations of forest trees plays an important role in forest tree 

improvement, conservation genetics, and containment of genetically modified trees (GM trees). Although trees are a very 

diverse group of woody plants, they share many common life 

traits that affect gene flow, which we discuss in this review. 

Some unfavorable processes in current forest tree ecosystems, 

such as habitat loss and fragmentation, increased environmental stress due to global climate change, introgression from domesticated trees into their wild relatives, introduction of maladapted germplasm during reforestation, etc., may badly affect 

naturally established balance between gene flow, isolation, and 

local adaptation. Therefore, estimation of gene flow becomes 

increasingly important for monitoring these processes and 

developing the best possible strategy to manage and protect 

forest ecosystems. Here, we review what has been done in this 

area recently and the methods and approaches currently used 

to measure gene flow in forest tree populations.

Link: http://dx.doi.org/10.1007/978-1-4614-0920-5_4

447. La Sorte FA, Jetz W. 2010. Avian distributions 

under climate change: towards improved projections. Journal of 

Experimental Biology 213:862-869.

Type: Journal 

Geographic Area: Global

Keywords: birds, biophysical model, bioclimate envelopment 

model, climate change, community composition, geographic 

range shifts, global climate change, migration

Abstract: Birds are responding to recent climate change in a 

variety of ways including shifting their geographic ranges to 

cooler climates. There is evidence that northern-temperate birds 

have shifted their breeding and non-breeding ranges to higher 

latitudes, and tropical birds have shifted their breeding ranges 

to higher altitudes. There is further evidence these shifts have 

affected migration strategies and the composition and structure 

of communities. Projections based on correlative distributional 

models suggest many birds will experience substantial pressures under climate change, resulting in range contraction and 

shifts. Inherent limitations of correlative models, however, 

make it difficult to develop reliable projections and detailed 

inference. Incorporating a mechanistic perspective into species 

distribution models enriches the quality of model inferences 

but also severely narrows the taxonomic and geographic relevance. Mechanistic distributional models have seen increased 

applications, but so far primarily in ectotherms. We argue that 

further development of similar models in birds would complement existing empirical knowledge and theoretical projections. 

The considerable data already available on birds offer an exciting basis. In particular, information compiled on flight performance and thermal associations across life history stages could 

be linked to distributional limits and dispersal abilities, which 

could be used to develop more robust and detailed projections. Yet, only a broadening of taxonomic scale, specifically 

CHAPTER 3 • MIGRATION • RESEARCH
160 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Geographic Area: China

Keywords: extreme weather event, hydropower projects, 

managed relocation, out-of-range translocation, population 

dynamics, tropical forest

Abstract: Assisted colonization of endangered species to 

locations outside their native ranges in response to projected 

climate change has emerged as a potential, but highly controversial conservation tool. The debate has been largely philosophical and hypothetical as little biological data exist in the 

literature. In 2006, nearly 1,000 endangered wild orchid plants 

belonging to 29 species were translocated to higher elevations 

in subtropical southwestern China in response to inundation 

threats from a hydropower project. We took advantage of this 

upward translocation to address one of the main biological 

concerns associated with assisted colonization, i.e., whether 

the target endangered species can survive in the novel environment that is projected to be suitable for them, sometime in the 

near future. We assessed the impacts of two extreme weather 

events, translocation shock and herbivory, on survival of 20 of 

these species and 462 individuals that were translocated beyond their current range vs. within that range. A cold spell in 

2008 on average caused 10% mortality, less than the mortality rate from herbivores. However, the cold spell was the only 

forces that extirpated an out-of-range population. No mortality resulted from a drought event in 2010. The 5-year survival 

percentages were not different between low and wide elevation 

species (69.3% mean ± 36.3% standard deviation vs. 67.3% 

± 30.9%). Orchids represent 10% of flowering plant diversity 

and are among the most endangered group of organisms due 

to combination of their often specialized ecological requirements, habitat destruction, and overexploitation. The demonstrated ability to survive extreme environmental challenges 

indicates that assisted colonization may be a viable conservation tool for the many endangered orchids worldwide due to 

climate change and/or other reasons. 

Link: http://english.xtbg.cas.cn/at/sr/201207/

P020120716522880199452.pdf

452. Lo Y, Blanco JA, Kimmins JP, Seely B, Welham 

C. 2011. Linking climate change and forest ecophysiology to 

project future trends in tree growth: a review of forest models.

Chapter 4. In: Blanco J, editor. Climate Change - Research and 

Technology for Adaptation and Mitigation. Rijeka, Croatia: 

InTech. p 63-87. 

Type: Book Section

Geographic Area: Western Canada

Compilers’ Keywords: species distribution, historical ecosystem, range shift

Introduction: Climate change is already altering tree species ranges, with tree lines shifting upwards and polewards 

around the world. A recent analysis of the potential effects 

of climate change on tree distribution in British Columbia 

By comparing the altitudinal distribution of 171 forest plant 

species between 1905 and 1985 and 1986 and 2005 along the 

entire elevation range (0 to 2600 meters above sea level) in 

west Europe, we show that climate warming has resulted in a 

significant upward shift in species optimum elevation averaging 29 meters per decade. The shift is larger for species restricted to mountain habitats and for grassy species, which are 

characterized by faster population turnover. Our study shows 

that climate change affects the spatial core of the distributional 

range of plant species, in addition to their distributional margins, as previously reported.

Link: http://www.researchgate.net/publication/224903439_A_

significant_upward_shift_in_plant_species_optimum_elevation_during_the_20th_century/file/9fcfd504cba44ab9ed.pdf

450. Linhart YB, Grant MC. 1996. Evolutionary significance of local genetic differentiation in plants. Annual Review 

of Ecology and Systematics 27:237-277.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: selection, genetic variation, microdifferentiation, evolutionary dynamics, natural selection

Abstract: The study of natural plant populations has provided 

some of the strongest and most convincing cases of the operation of natural selection currently known, partly because 

of amenability to reciprocal transplant experiments, common 

garden work, and long-term in situ manipulation. Genetic differentiation among plant populations over small scales (a few 

cm to a few hundred cm) has been documented and is reviewed 

here, in herbaceous annuals and perennials, woody perennials, 

aquatics, terrestrials, narrow endemics, and widely distributed species. Character differentiation has been documented 

for most important features of plant structure and function. 

Examples are known for seed characters, leaf traits, phenology, physiological and biochemical activities, heavy metal 

tolerance, herbicide resistance, parasite resistance, competitive ability, organellar characters, breeding systems, and life 

history. Among the forces that have shaped these patterns of 

differentiation are toxic soils, fertilizers, mowing and grazing, 

soil moisture, temperature, light intensity, pollinating vectors, 

parasitism, gene flow, and natural dynamics. The breadth and 

depth of the evidence reviewed here strongly support the idea 

that natural selection is the principal force shaping genetic architecture in natural plant populations; that view needs to be 

more widely appreciated than it is at present.

451. Liu H, Feng C-L, Chen B-S, Wang Z-S, Xie X-Q, 

Deng Z-H, Wei X-L, Liu S-Y, Zhang Z-B, Luo Y-B. 

2012. Overcoming extreme weather challenges: successful but 

variable assisted colonization of wild orchids in southwestern 

China. Biological Conservation 150:68-75.

Type: Journal 

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 161

Abstract: The ranges of plants and animals are moving in 

response to recent changes in climate. As temperatures rise, 

ecosystems with ‘nowhere to go’, such as mountains, are considered to be more threatened. However, species survival may 

depend as much on keeping pace with moving climates as the 

climate’s ultimate persistence. Here we present a new index of 

the velocity of temperature change (km yr-1), derived from spatial gradients (°C km-1) and multimodel ensemble forecasts of 

rates of temperature increase (°C yr-1) in the twenty-first century. This index represents the instantaneous local velocity along 

Earth’s surface needed to maintain constant temperatures, and 

has a global mean of 0.42 km yr-1 (A1B emission scenario). 

Owing to topographic effects, the velocity of temperature 

change is lowest in mountainous biomes such as tropical and 

subtropical coniferous forests (0.08 km yr-1), temperate coniferous forest, and montane grasslands. Velocities are highest in 

flooded grasslands (1.26 km yr-1), mangroves and deserts. High 

velocities suggest that the climates of only 8% of global protected areas have residence times exceeding 100 years. Small 

protected areas exacerbate the problem in Mediterranean-type 

and temperate coniferous forest biomes. Large protected areas 

may mitigate the problem in desert biomes. These results indicate management strategies for minimizing biodiversity loss 

from climate change. Montane landscapes may effectively 

shelter many species into the next century. Elsewhere, reduced 

emissions, a much expanded network of protected areas, or efforts to increase species movement may be necessary.

Link: http://www.ncbi.nlm.nih.gov/pubmed/20033047

454. Lu P, Man R. 2011. Assessment of assisted migration 

effects on spring bud flush in white spruce (Picea glauca) seedlings. The Forestry Chronicle 87:391-397.

Type: Journal 

Geographic Area: Ontario, Canada

Keywords: bud flush, provenance, frost damage

Abstract: In a changing climate, delaying the time of bud flush 

may be advantageous to boreal forest species to reduce the risk 

of spring frost damage. In this study, we examined the potential effect of assisted migration of tree seed on time to bud flush 

for white spruce. Flushing times of seedlings from 23 white 

spruce provenances from Ontario were observed under varying temperature conditions simulated in controlled environment chambers. Results indicated that time to bud flush varied 

considerably among provenances. Although higher temperatures significantly promoted bud flushing for all provenances, 

provenance-by-temperature interactions were negligible, indicating stable performance of white spruce provenances for this 

adaptive trait. Spatial patterns of variation among provenances 

in bud flushing were not consistent with patterns found in 

range-wide provenance tests. Assisted migration of tree seed 

across relatively short distances is unlikely to delay bud flushing time in white spruce. Some southern populations may flush 

suggested that important timber species including white 

spruce and lodgepole pine may lose suitable habitat and suffer adversely from a combination of warming trends and reduced growing season precipitation. In contrast, species such 

as Douglas fir and ponderosa pine may actually expand their 

range and potentially show improved growth rates in parts 

of their existing range. A recent study in the mountains of interior British Columbia showed how at high elevation, trees 

historically responded positively to increased temperatures, 

while at low elevations trees showed a negative response 

to growing season maximum temperature and a positive 

correlation with growing season precipitation. Given these 

species-specific responses it is not surprising that recent 

research has failed to identify direct links between warmer 

temperatures and observed changes in species ranges. The 

important ecological and socio-economic consequences of 

such changes have prompted multiple modelling efforts to 

predict the future location of habitat suitable for tree species and to assess the potential implications for tree growth 

of changes in climate. Defining such areas and estimating 

the losses or gains due to climate change in timber production have important consequences on forest management 

and conservation. The most popular approaches to project 

future areas of suitable habitat for commercial tree species 

have involved analysis of historical records of tree lines 

in boreal and alpine environments, using climate envelope 

models. Similarly, dendroclimatology (studying historical 

tree growth rates by analyzing tree ring width) has been used 

to link climate and tree growth rates. These approaches are 

based mostly on climatic information, although their combination with other information such as soil or topography 

has been used to produce maps of potential future habitat 

suitability. Such predictions are useful to understand the relationships between climate and tree distribution, abundance 

and growth, and could be a starting point for helping to plan 

forest management at broad scales under changing climate. 

However, such approach has several shortcomings, which 

have been discussed in the scientific literature before but it 

seems that this discussion has not been translated into the 

forest management community yet. 

Link: http://www.researchgate.net/publication/221915588_

Linking_Climate_Change_and_Forest_Ecophysiology_to_

Project_Future_Trends_in_Tree_Growth_A_Review_of_Forest_Models/file/60b7d5141a7b103241.pdf

453. Loarie SR, Duffy PB, Hamilton H, Asner GP, 

Field CB, Ackerly DD. 2009. The velocity of climate 

change. Nature 462:1052-1055.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: animals, biodiversity, conservation, 

ecosystem, global warming, biological models, temporal factors, migration, species distribution

CHAPTER 3 • MIGRATION • RESEARCH
162 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

discussed, including intrinsic migration capabilities, barriers to 

migration, the role of outlier populations in increasing migration rates, the role of climate in setting range limits and variation in species range sizes.

Link: http://citeseerx.ist.psu.edu/viewdoc/download?doi=10.1

.1.215.1388&rep=rep1&type=pdf

456. Malcolm JR, Liu C, Neilson RP, Hansen L, 

Hannah L. 2006. Global warming and extinctions of endemic species from biodiversity hotspots. Conservation Biology 

20:538-548.

Type: Journal 

Geographic Area: Global

Keywords: biomes, climate change, general circulation models, global vegetation models, migration, species extinctions

Abstract: Global warming is a key threat to biodiversity, but 

few researchers have assessed the magnitude of this threat at the 

global scale. We used major vegetation types (biomes) as proxies for natural habitats and, based on projected future biome 

distributions under doubled-CO2 climates, calculated changes 

in habitat areas and associated extinctions of endemic plant and 

vertebrate species in biodiversity hotspots. Because of numerous uncertainties in this approach, we undertook a sensitivity 

analysis of multiple factors that included (1) two global vegetation models, (2) different numbers of biome classes in our 

biome classification schemes, (3) different assumptions about 

whether species distributions were biome specific or not, and 

(4) different migration capabilities. Extinctions were calculated using both species-area and endemic-area relationships. 

In addition, average required migration rates were calculated 

for each hotspot assuming a doubled-CO2 climate in 100 years. 

Projected percent extinctions ranged from < 1 to 43% of the 

endemic biota (average 11.6%), with biome specificity having 

the greatest influence on the estimates, followed by the global 

vegetation model and then by migration and biome classification assumptions. Bootstrap comparisons indicated that effects 

on hotpots as a group were not significantly different from 

effects on random same biome collections of grid cells with 

respect to biome change or migration rates; in some scenarios, 

however, hotspots exhibited relatively high biome change and 

low migration rates. Especially vulnerable hotspots were the 

Cape Floristic Region, Caribbean, Indo-Burma, Mediterranean 

Basin, Southwest Australia, and Tropical Andes, where plant 

extinctions per hotspot sometimes exceeded 2,000 species. 

Under the assumption that projected habitat changes were attained in 100 years, estimated global-warming-induced rates 

of species extinctions in tropical hotspots in some cases exceeded those due to deforestation, supporting suggestions 

that global warming is one of the most serious threats to the 

planet’s biodiversity.

Link: http://www.atlanticcoastconservancy.org/Global%20

Warming%20Articles/Malcolm%20et%20al.%202005.pdf

earlier at more northerly sites, which would exacerbate spring 

frost risk. Tree improvement may be an effective approach to 

utilize among- and within provenance variation to enhance this 

fitness trait for better climatic adaptation.

455. Malcolm JR, Markham A, Neilson RP, 

Garaci M. 2002. Estimated migration rates under 

scenarios of global climate change. Journal of Biogeography 

29:835-849.

Type: Journal 

Geographic Area: North America

Keywords: global warming, plant migration, biomes, greenhouse effect, biodiversity

Abstract: Greenhouse-induced warming and resulting shifts 

in climatic zones may exceed the migration capabilities of 

some species. We used fourteen combinations of General 

Circulation Models (GCMs) and Global Vegetation Models 

(GVMs) to investigate possible migration rates required under 

CO2-doubled climatic forcing. Migration distances were calculated between grid cells of future biome type x and nearest 

same-biome-type cells in the current climate. In ‘base-case’ 

calculations, we assumed that 2·CO2 climate forcing would 

occur in 100 years, we used ten biome types and we measured 

migration distances as straight-line distances ignoring water 

barriers and human development. In sensitivity analyses, we 

investigated different time periods of 2·CO2 climate forcing, 

more narrowly defined biomes and barriers because of water 

bodies and human development. In the base-case calculations, 

average migration rates varied significantly according to the 

GVM used (BIOME3 vs. MAPSS), the age of the GCM (older- vs. newer-generation GCMs), and whether or not GCMs 

included sulfate cooling or CO2 fertilization effects. However, 

high migration rates (≥ 1000 m year-1) were relatively common in all models, consisting on average of 17% grid cells for 

BIOME3 and 21% for MAPSS. Migration rates were much 

higher in boreal and temperate biomes than in tropical biomes. 

Doubling of the time period of 2·CO2 forcing reduced these 

areas of high migration rates to c. 12% of grid cells for both 

BIOME3 and MAPSS. However, to obtain migration rates 

in the Boreal biome that were similar in magnitude to those 

observed for spruce when it followed the retreating North 

American Glacier, a radical increase in the period of warming was required, from 100 to > 1,000 years. A reduction in 

biome area by an order of magnitude increased migration rates 

by one to three orders of magnitude, depending on the GVM. 

Large water bodies and human development had regionally 

important effects in increasing migration rates. In conclusion, 

evidence from coupled GCMs and GVMs suggests that global 

warming may require migration rates much faster than those 

observed during postglacial times and hence has the potential 

to reduce biodiversity by selecting for highly mobile and opportunistic species. Several poorly understood factors that are 

expected to influence the magnitude of any such reduction are 

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 163

to policymakers just about in time for him to retire. In a world 

in which many scientists fret about the toll that global warming is exacting on nature, O’Neill is actually doing something 

about it. A research scientist for the British Columbia Ministry 

of Forests, he is moving seedlings to areas that are outside their 

current comfort zone to test how they might handle the warmer 

conditions of the future. And he is behind a government push 

to move tree populations into new areas to prepare them for the 

warming climate.

459. Maschinski J, Haskins KE. 2012. Plant reintroduction in a changing climate: promises and perils. Washington 

(DC): Island Press. 402 p. 

Type: Book

Geographic Area: North America

Compilers’ Keywords: migration, relocation, review, guidelines, plants, conservation, species extinction

Introduction: In the face of mounting numbers of plant species at risk of extinction, increasing rates of habitat destruction, 

spreading invasive species, and effects of climate change, there 

is a great need for urgent action to preserve species before they 

are extirpated. This book tells a story of the good fight to save 

and restore some of the rarest plant species in the world. A 

review of plant reintroductions is paramount because plant 

management techniques are fundamentally different from 

those used for animals. Herein we take stock of our progress 

with reintroduction in an effort to facilitate the wise decisions 

needed to preserve future biodiversity. This book is intended 

to be a resource for students, practitioners, and conservation 

scientists.

Link: http://islandpress.org/ip/books/book/islandpress/P/

bo8073733.html

460. Maschinski J, Baggs JE, Quintana-Ascencio 

PF, Menges ES. 2006. Using population viability analysis 

to predict the effects of climate change on the extinction risk 

of an endangered limestone endemic shrub, Arizona cliffrose.

Conservation Biology 20:218-228.

Type: Journal 

Geographic Area: Arizona, USA

Keywords: drought, endangered species, global warming, 

population viability analysis, Purshia subintegra

Abstract: The threat of global warming to rare species is a 

growing concern, yet few studies have predicted its effects on 

rare populations. Using demographic data gathered in both 

drought and nondrought years between 1996–2003 in central 

Arizona upper Sonoran Desert, we modeled population viability for the federally endangered Purshia subintegra (Kearney) 

Henrickson (Arizona cliffrose). We used deterministic matrix 

projection models and stochastic models simulating weather 

conditions during our study, given historical weather variation 

457. Mandák B, Placková I. 2009. How does population genetic diversity change over time? An experimental 

seed bank study of Atriplex tatarica (Chenopodiaceae). Flora 

- Morphology, Distribution, Functional Ecology of Plants 

204:423-433.

Type: Journal 

Geographic Area: Czech Republic, Europe

Keywords: allozyme, competition, heterozygosity, seed bank, 

succession, Wright’s F statistics

Abstract: Atriplex tatarica is an annual, early successional, 

facultative halophilic species of frequently disturbed humanmade habitats in Central and Eastern Europe. We investigated 

to what extent the plants grown from seeds extracted from soil 

seedbank differed genetically to mature aboveground plants 

in experimental populations of A. tatarica over two successive years. At each of five plots 50 aboveground plants and 

50 plants extracted from seeds stored in soil were assayed for 

allozyme analysis in 2003 and 2004. At the start of experiment, 

we introduced 1,000 seeds of the study species into each of five 

experimental plots. While the species dominated in all of the 

experimental plots in the first year, the second year A. tatarica

cover age decreased dramatically. Over all allele frequencies 

of soil seeds and mature plants showed significant differences 

between life history stages in both years, but not within years in 

soil seeds as well as mature plants stages. While mature plants 

showed a significantly greater amount of single and multilocus 

heterozygosity in both consecutive years, comparison between 

years did not yield any significant differences. In the same 

way, despite a relatively large seedbank the species population 

genetic parameters, i.e., allelic richness (A), observed heterozygosity (Ho), gene diversity (Hs), inbreeding coefficient (FIS) 

and fixation index (FST), did not change over the years between 

as well as within life history stages. The soil seeds and mature 

plants significantly differed in Ho, Hs and FIS, while the A and 

FST were not significantly different between life history stages.

458. Marris E. 2009. Planting the forest for the future.

Nature 459:906-908.

Type: Journal 

Geographic Area: North America

Compilers’ Keywords: assisted migration, forest plant species, conservation, climate change, common garden, plant migration, Assisted Migration Adaptation Trial

Introduction: A valley in British Columbia, a few kilometres 

outside the town of Vernon, orderly rows of trees run alongside 

the road. Many of the conifers stand tall and full, producing 

seeds destined for plantations around the province. But one 

dusty brown field is filled with lines of seedlings just ankle 

high. Greg O’Neill, 45, who planted the trees in April, walks 

among them with a slightly paternal air. They are part of a very 

slow experiment that will yield the first results that are useful 

CHAPTER 3 • MIGRATION • RESEARCH
164 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

indicate that natural genealogical lineages of cultivars, such as 

Garnet mountain brome, have dispersed and succeeded over 

broad geographical regions. However, more research and plant 

material work are needed before specific recommendations 

can be made over the entire species distribution.

Link: http://nativeplantnetwork.org/Content/Articles/6-

1NPJ29-35.pdf

462. Matthews SN, Iverson LR, Prasad AM, 

Peters MP, Rodewald PG. 2011. Modifying climate 

change habitat models using tree species-specific assessments of 

model uncertainty and life history-factors. Forest Ecology and 

Management 262:1460-1472.

Type: Journal 

Geographic Area: Eastern USA

Keywords: climate change adaptation, Eastern United 

States, trees, species distribution models, disturbance, model 

uncertainty

Abstract: Species distribution models (SDMs) to evaluate 

trees’ potential responses to climate change are essential for developing appropriate forest management strategies. However, 

there is a great need to better understand these models’ limitations and evaluate their uncertainties. We have previously 

developed statistical models of suitable habitat, based on both 

species’ range and abundance, to better understand potential 

changes of 134 tree species habitats in the eastern United States 

(http://www.nrs.fs.fed.us/atlas). Our focus here is to build on 

these results via a more robust assessment framework called 

modification factors (ModFacs) that is made up of five components. ModFac 1 addresses nine biological characteristics (e.g., 

shade tolerance and seedling establishment) that quantify the 

influence of species life-history traits. ModFac 2 considers 12 

disturbance characteristics (e.g., insect pests, drought, and fire 

topkill) which address the capacity of a species to tolerate and 

respond to climate-induced changes in habitat. ModFac 3–5 

distill the tree SDM results and facilitate communication of 

model uncertainty; we quantified the variability in projected 

change for General Circulation Models (GCM) and emissions 

scenarios (ModFac 3), the extent to which each species’ habitat 

intersects novel climate conditions (Mod-Fac4), and accounted for long-distance extrapolations beyond a species’ current 

range (ModFac5). The life-history components of ModFacs 

1 and 2 demonstrate the marked variability among species in 

terms of biological and disturbance characteristics, suggesting diverse abilities to adapt to climate change. ModFacs 3–5 

show that the information from the SDMs can be enhanced by 

quantifying the variability associated with specific GCM/emission scenarios, the emergence of novel climates for particular 

tree species, and the distances of species habitat shifts with climate change. The ModFacs framework has high interpretive 

value when considered in conjunction with the outputs of species habitat models for this century. Importantly, the intention 

and under scenarios of increased aridity. Our models suggest 

that the P. subintegra population in Verde Valley is slowly declining and will be at greater risk of extinction with increased 

aridity. Across patches at a fine spatial scale, demographic 

performance was associated with environmental factors. Moist 

sites (patches with the highest soil moisture, lowest sand content, and most northern aspects) had the highest densities, highest seedling recruitment, and highest risk of extinction over 

the shortest time span. Extinction risk in moist sites was exacerbated by higher variance in recruitment and mortality. Dry 

sites had higher cumulative adult survival and lower extinction 

risk but negative growth rates. Steps necessary for the conservation of the species include introductions at more northern 

latitudes and in situ manipulations to enhance seedling recruitment and plant survival. We demonstrate that fine spatial-scale 

modeling is necessary to predict where patches with highest 

extinction risk or potential refugia for rare species may occur. 

Because current climate projections for the 21st century imply 

range shifts at rates of 300 to 500 km/century, which are beyond even exceptional examples of shifts in the fossil record of 

100–150 km, it is likely that preservation of many rare species 

will require human intervention and a long-term commitment. 

Global warming conditions are likely to reduce the carrying 

capacity of many rare species’ habitats.

Link: http://pascencio.cos.ucf.edu/classes/Research%20publications/PDFManuscripts/Maschinski%20et%20al%202006.

pdf

461. Massa AN, Larson SR. 2005. Phylogeography 

of North American mountain bromes. Native Plants Journal 

6:29-35.

Type: Journal 

Geographic Area: North America

Keywords: AFLP, genetic diversity, Bromus carinatus

Abstract: Although native grasses are often desired and used 

for revegetation of disturbed areas, genetic differences may exist within and among natural and cultivated germplasm sources. 

This phylogeographic study compares geographic origin and 

genealogical linkages of 25 natural and cultivated germplasm 

sources of mountain brome (Bromus carinatus Hook. & Arn. 

[Poaceae]) from western North America. Significant variation 

among accessions (FST = 0.70) was detected by analysis of molecular variance (AMOVA), based on the number of amplified 

fragment length polymorphisms (AFLPs) between individual 

plants. Likewise, significant differences among 4 hierarchical 

genotypic groups, encompassing all but 5 unique accessions, 

were also detected (FCT = 0.47).This study identified at least 

one well-defined genealogical lineage, comprising 8 accessions, distributed over a broad geographic region and different 

ecosystems of western North America. Two other hierarchical 

groups, comprising 6 accessions and 3 accessions, were located within or near specific ecoregions. Results of this study 

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 165

while the needs of conservation practice have had little influence on seed dispersal research. Yet seed dispersal interacts 

decisively with the major drivers of biodiversity change in 

the 21st century: habitat fragmentation, overharvesting, biological invasions, and climate change. We synthesize current 

knowledge of the effects these drivers have on seed dispersal 

to identify research gaps and to show how this information 

can be used to improve conservation management. The drivers, either individually, or in combination, have changed the 

quantity, species composition, and spatial pattern of dispersed 

seeds in the majority of ecosystems worldwide, with inevitable 

consequences for species survival in a rapidly changing world. 

The natural history of seed dispersal is now well understood in 

a range of landscapes worldwide. Only a few generalizations 

that have emerged are directly applicable to conservation management, however, because they are frequently confounded by 

site-specific and species-specific variation. Potentially synergistic interactions between disturbances are likely to exacerbate the negative impacts, but these are rarely investigated. We 

recommend that the conservation status of functionally unique 

dispersers be revised and that the conservation target for key 

seed dispersers should be a population size that maintains their 

ecological function, rather than merely the minimum viable 

population. Based on our analysis of conservation needs, seed 

dispersal research should be carried out at larger spatial scales 

in heterogeneous landscapes, examining the simultaneous impacts of multiple drivers on community-wide seed dispersal 

networks.

Link: http://www.researchgate.net/publication/235331752_Seed_dispersal_in_changing_landscapes/

file/9fcfd510e8cfabead7.pdf

465. McDonald-Madden E, Runge MC, Possingham 

HP, Martin TG. 2011. Optimal timing for managed relocation of species faced with climate change. Nature Climate 

Change 1:261-265.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: assisted migration, decision framework, management, model

Abstract: Managed relocation is a controversial climate-adaptation strategy to combat negative climate change impacts on 

biodiversity. While the scientific community debates the merits 

of managed relocation, species are already being moved to new 

areas predicted to be more suitable under climate change. To inform these moves, we construct a quantitative decision framework to evaluate the timing of relocation in the face of climate 

change. We find that the optimal timing depends on many factors, including the size of the population, the demographic costs 

of translocation and the expected carrying capacities over time 

in the source and destination habitats. In some settings, such 

as when a small population would benefit from time to grow 

of this assessment was not to create a static scoring system, but 

to broadly assess species characteristics that likely will play 

an important role in adaptation to climate change. We believe 

these scores based on biological, disturbance, and model synthesis factors provide an important expansion of interpretive 

and practical value to habitat model projections.

Link: http://obcp.osu.edu/files/Rodewald%201.pdf

463. McCarragher SR, Goldblum D, Rigg LS. 2011.

Geographic variation of germination, growth, and mortality in 

sugar maple (Acer saccharum): common garden and reciprocal 

dispersal experiments. Physical Geography 32:1-21.

Type: Journal 

Geographic Area: Northeastern USA

Compilers’ Keywords: biogeography, GCM, migration, 

conservation

Abstract: If current temperature trends continue and increase 

as predicted by general circulation models, the persistence 

and migration of sugar maple (Acer saccharum) at its northern 

limit will become crucial to its continued existence in North 

America. Specifically, anthropogenic warming may affect germination success and growth of sugar maple. To evaluate this 

potential limitation, our study addresses two questions. The 

first involves a growth chamber: Do temperatures at and above 

(1, 7, and 14 °C) sugar maple’s optimal germination temperature affect its germination? The second involves a common 

garden: Do sugar maple germination and subsequent growth 

and mortality rates show evidence of geographic variation 

under current climatic conditions? While sugar maple germinated successfully at 7 °C in the lab, field results suggest that 

other environmental variables may limit its future establishment as climate changes. Germination of seeds from sugar 

maple’s northern range limit was significantly reduced under 

warmer conditions, and subsequent seedling mortality rates 

were significantly increased when grown at a more southerly 

latitude. Local adaptation was evident with respect to germination, survival, and growth in the field. Ultimately, results from 

this study further the understanding of how predicted anthropogenic climate change may affect the regeneration of sugar 

maple in the future.

464. McConkey KR, Prasad S, Corlett RT, CamposArceiz A, Brodie JF, Rogers H, Santamaria L. 2012.

Seed dispersal in changing landscapes. Biological Conservation 

146:1-13.

Type: Journal 

Geographic Area: Global

Keywords: biological invasions, climate change, fragmentation, hunting, overharvesting, seed dispersal

Abstract: A growing understanding of the ecology of seed dispersal has so far had little influence on conservation practice, 

CHAPTER 3 • MIGRATION • RESEARCH 
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records suggest that tree populations are capable of rapid migration when climate warms. Fossil pollen is commonly interpreted as suggesting that the range of many temperate tree 

species expanded at rates of 100–1,000 m/yr during the early 

Holocene. We used chloroplast DNA surveys to show that 

the geography of postglacial range expansion in two eastern 

North American tree species differs from that expected from 

pollen-based reconstructions and from patterns emerging from 

European molecular studies. Molecular evidence suggests that 

American beech (Fagus grandifolia) and red maple (Acer rubrum) persisted during the late glaciation as low-density populations, perhaps within 500 km of the Laurentide Ice Sheet. 

Because populations were closer to modern range limits than 

previously thought, postglacial migration rates may have been 

slower than those inferred from fossil pollen. Our estimated 

rates of < 100 m/yr are consistent with model predictions based 

on life history and dispersal data, and suggest that past migration rates were substantially slower than the rates that will be 

needed to track 21st-century warming.

Link: http://forestbiology.ucdavis.edu/docs/MolecularIndicators_treeMigration.pdf

468. McLane SC. 2011. Establishment and growth responses of whitebark and lodgepole pine populations in a 

changing climate [Dissertation]. Vancouver, British Columbia, 

Canada: The University of British Columbia. 125 p.

Type: Dissertation

Geographic Area: North America

Compilers’ Keywords: Pinus contorta ssp. latifolia, Pinus 

albicaulis, climate change, common garden, migration

Abstract: Climate change will affect the regeneration, growth, 

survival and distribution of trees. Here, I use common gardens 

to empirically test establishment, growth and the potential for 

persistence, adaptation and migration for two iconic North 

American trees, whitebark pine (Pinus albicaulis) and lodgepole 

pine (Pinus contorta ssp. latifolia). Whitebark pine is of conservation concern due to range-wide diebacks, while lodgepole 

pine is critical to forest productivity and carbon sequestration. 

Whitebark seeds were planted north of the current range in areas 

predicted to be climatically suitable through the 2050s; these 

germinated and survived in varying proportions at all locations. 

Establishment and growth were positively affected by moderate 

snow-cover durations, heavier seed weights, and warmer provenance temperatures. Whitebark pine seedlings grown from 

seeds sown in growth chambers spanning current and predictedfuture temperatures demonstrated positive responses to warmer 

growing seasons. Lodgepole pine seedlings in the same chambers outgrew the whitebark pine seedlings at all but the coldest temperatures. Together, these results suggest that whitebark 

pine may lose its competitive advantage to other species within 

its narrow alpine-treeline niche as the climate warms, but that it 

is capable of establishing in climatically-suitable areas north of 

before risking translocation losses, haste is ill advised. We also 

find that active adaptive management is valuable when the effect of climate change on source habitat is uncertain, and leads 

to delayed movement. 

Link: http://www.seaturtle.org/PDF/McDonald-MaddenE_2011_NatureClimChange.pdf

466. McKenney DW, Pedlar J, O’Neill GA. 2009.

Climate change and forest seed zones: past trends, future 

prospects and challenges to ponder. The Forestry Chronicle 

85:258-265.

Type: Journal 

Geographic Area: Canada

Keywords: climate change, seed sources, British Columbia, 

Ontario, Douglas-fir, seed transfer, assisted migration

Abstract: Canada regenerates more than 400,000 ha of forest 

land annually through planting and seeding operations. Much 

of the stock for this effort is selected to be climatically suited to 

the planting site—a match that is often facilitated through the 

development of seed zones. However, if climate change proceeds as predicted, stock that is well matched under current climate will be growing in sub-optimal conditions within the next 

20 to 50 years—in some parts of the country, trees may already 

be growing outside their optimal climates. To provide a sense 

of the magnitude of these changes, we present past and predicted future climate trends for Ontario and British Columbia 

seed zones. For Ontario, over the period 1950 to 2005, minimum temperature of the coldest month has already increased 

by up to 4.3 °C, growing season has lengthened by up to 6 days, 

and precipitation during the growing season has increased by 

up to 26%. Changes were more pronounced across British 

Columbia’s Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco) 

seed zones, with minimum temperature increasing by up to 8 

°C, a growing season extension of up to 30 days, and growing season precipitation increases of up to 40%. Projections for 

the end of the current century include: minimum temperature 

increase of 5 °C to 10 °C, growing season extension of 31 to 60 

days, and growing season precipitation increases of 3% to 42% 

across the seed zones in both provinces. These changes are certain to have extensive impacts on forest ecosystems. We briefly 

discuss 3 forest management adaptation strategies intended to 

mitigate the negative impacts of climate change in Canada.

467. McLachlan JS, Clark JS, Manos PS. 2005.

Molecular indicators of tree migration capacity under rapid climate change. Ecology 86:2088-2098.

Type: Journal 

Geographic Area: North America, Europe

Keywords: climate change, molecular markers, paleoecology, 

range expansion

Abstract: Recent models and analyses of paleoecological 

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Our experiment provides a preliminary test of models predicting the existence of climatically suitable whitebark pine habitat 

north of the current species ranges. More broadly, our techniques and results inform the development of scientific guidelines for assisting the migration of other species that are highly 

threatened by climate change. Applied case studies of this kind 

are critical for assessing the utility of species distribution models as conservation planning tools.

470. Menges ES. 2008. Restoration demography and genetics of plants: when is a translocation successful? Australian 

Journal of Botany 56:187-196.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: assisted migration, key species, 

review

Abstract: Restorations are complex, often involving restoring ecological processes, vegetation structure, and species’ 

populations. One component of restorations is translocation 

of key species. Translocations (introductions, reintroductions, 

augmentations) are often necessary to recover species diversity and install key species. In this review, I consider the ways 

translocations have been evaluated at various stages during the 

process of restoration. Vital rates (survival, growth, fecundity) 

of propagules (seeds, transplants) are commonly used to evaluate initial success. Transplants usually provide greater initial 

success than do sown seeds. Beyond initial rates, completion 

of the life cycle through flowering, fruiting, dispersal and subsequent seedling recruitment is a key benchmark. Modelling 

population viability of translocated populations is a logical 

next step and can bring in many powerful inferential tools. Of 

factors affecting the success of translocations, genetic issues 

are paramount, as restorationists need to consider inbreeding 

depression, reproductive viability, local adaptation, and evolutionary potential of translocated populations. The success of 

translocations is also clearly context-dependent, with herbivory, disturbance, competition and other ecological factors important. Future translocations need to make better use of comparisons with reference populations, a long-term perspective 

on success and an experimental framework that can provide 

both practical and basic knowledge. Demographic data collection and analysis in restorations has great potential to elucidate 

causes of translocation failure and improve the prognosis of 

future restorations.

Link: http://pascencio.cos.ucf.edu/classes/Restoration%20

Ecology/Menges%202008%20restoration%20demography.pdf

471. Millar MA, Byrne M, Nuberg IK, Sedgley M. 

2012. High levels of genetic contamination in remnant populations of Acacia saligna from a genetically divergent planted 

stand. Restoration Ecology 20:260-267.

Type: Journal 

its current range. Using tree-ring data from long-term lodgepole 

pine common garden trials, I built universal growth-trend response functions to forecast future growth trends relative to genetics, climate and tree age. The models predict growth reductions for all populations by the end of the 21st century based on 

middle-of-the-road climate models, except in far northern areas 

near and within Yukon, Canada. Analogous models built using 

summer and winter climate indices indicate that the growth 

declines are primarily caused by warmer summers, and may 

be offset by growth increases resulting from warmer winters. 

I found that populations are most sensitive to annual temperatures and summer aridity, but that sensitivity to climate varies 

due to local adaptation. Overall, my research will help forest 

professionals and conservationists forecast changes in forest 

productivity and species growth and survival under warming 

temperatures.

Link: https://circle.ubc.ca/bitstream/handle/2429/34087/

ubc_2011_spring_mclane_sierra.pdf

469. McLane SC, Aitken SN. 2012. Whitebark pine 

(Pinus albicaulis) assisted migration potential: testing establishment north of the species range. Ecological Applications 

22:142-153.

Type: Journal 

Geographic Area: Western Canada

Keywords: climate change, common garden, managed relocation, quantitative genetics, species distribution model, species range limits

Abstract: The translocation of species into habitable locations outside of their current ranges, termed assisted migration, 

has been proposed as a means of saving vulnerable species 

from extinction as a result of climate change. We explore the 

use of this controversial technique using a threatened keystone 

species in western North America, whitebark pine (Pinus albicaulis), as a case study. Species distribution models predict 

that whitebark pine will be extirpated from most of its current 

range as temperatures rise over the next 70 years. However, 

the same models indicate that a large area within northwestern 

British Columbia, Canada, is climatically suitable for the species under current conditions and will remain so throughout the 

21st century. To test the capacity of whitebark pine to establish 

relative to climatic and habitat features within its predicted climatic range, we planted seeds from seven populations in eight 

locations spanning from 600 km southeast to 800 km northwest of the northern boundary of the current species range. 

During the first three growing seasons, germination occurred 

in all locations. Nearly three times as many treated (induced 

maturation and broken dormancy) than untreated seeds germinated, and most treated seeds germinated a year earlier than the 

untreated seeds. Germination, survival, and growth were primarily influenced by seed mass, site climate conditions related 

to the duration of snow cover, and provenance temperature. 

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168 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

(Panthera tigris) appeared to represent assemblages that would 

not be conserved by ex situ host conservation. In contrast, for 

the cases of dependent species we examined involving a single 

dependent species (internal parasites of birds and the mite 

Geckobia naultina on Duvaucel’s gecko), ex situ conservation 

of the host species would also conserve the dependent species. 

However, moving dependent species with their hosts may be 

insufficient to maintain viable populations of the dependent 

species, and additional conservation strategies such as supplementing populations may be needed.

Link: http://www.ncbi.nlm.nih.gov/pubmed/22443127

473. Mooring JS. 2008. An Eriophyllum lanatum

(Asteraceae) hybrid zone in Oregon. Madroño 55:269-279.

Type: Journal 

Geographic Area: Oregon, USA

Keywords: artificial hybridizations, Asteraceae, barriers to interbreeding, Eriophyllum, hybrid zone, pollen fertility, species 

complex, supernumerary chromosomes

Abstract: Eriophyllum lanatum vars. achilleoides and leucophyllum intergrade in southwestern Oregon. Some populations cannot be unequivocally assigned to either variety. 

Chromosome counts showed polyploid populations predominating where the varieties meet in southwestern Oregon. 

Pollen fertility, estimated by cotton blue-lactophenol staining, 

was the main criterion used to assess barriers to interbreeding. 

Artificial hybridizations between diploids revealed barriers 

to interbreeding between vars. achilleoides and leucophyllum, 

and between each of them and a morphologically intermediate 

population. The barriers to interbreeding are substantially less 

developed than those present between most of the other eight 

varieties of the E. lanatum complex. Supernumerary chromosomes are postulated to be adaptive in Eriophyllum hybrid 

zones.

474. Moritz C, Patton JL, Conroy CJ, Parra JL, 

White GC, Beissinger SR. 2008. Impact of a century 

of climate change on small-mammal communities in Yosemite 

National Park, USA. Science 322:261-264.

Type: Journal 

Geographic Area: California, USA

Compilers’ Keywords: land-use change, modelling, migration, threatened species, high elevation

Abstract: We provide a century-scale view of small-mammal 

responses to global warming, without confounding effects 

of land-use change, by repeating Grinnell’s early–20th century survey across a 3,000-meter-elevation gradient that spans 

Yosemite National Park, California, USA. Using occupancy 

modeling to control for variation in detectability, we show substantial (~500 meters on average) upward changes in elevational limits for half of 28 species monitored, consistent with the 

Geographic Area: Australia

Keywords: fragmented vegetation, gene flow, genetic contamination, paternity analysis, pollen dispersal, risk assessment

Abstract: It is essential to understand the patterns of pollen 

dispersal in remnant vegetation occupying highly disturbed 

landscapes in order to provide sustainable management options and to inform restoration programs. Direct and indirect 

methods of paternity analysis were used to detect genetic contamination via inter-subspecific pollen dispersal from a planted 

stand of nonlocal Acacia saligna ssp. saligna (ms) into remnant 

roadside patches of local A. saligna ssp. lindleyi (ms). Genetic 

contamination was detected in 25.5% (indirect paternity assignment) to 32% (direct paternity assignment) of ssp. lindleyi

progeny and occurred over a distance of 1.6 km. The results 

support studies that suggest genetic continuity is maintained 

by high levels of pollen dispersal in temperate entomophilous 

species. The results also indicate that patchily distributed remnant populations may be exposed to substantial amounts of 

genetic contamination from large scale restoration with native 

taxa in the highly fragmented agricultural landscape of southern Western Australia. Management practices to reduce the 

risk of genetic contamination are considered.

472. Moir ML, Vesk PA, Brennan KE, Poulin R, 

Hughes L, Keith DA, McCarthy MA, Coates DJ. 

2012. Considering extinction of dependent species during 

translocation, ex situ conservation, and assisted migration of 

threatened hosts. Conservation Biology 26:199-207.

Type: Journal 

Geographic Area: Australia

Keywords: assisted colonization, coextinction conservation 

planning, extinction risk, introductions, managed relocation, 

parasites, plant-insect interactions, translocation

Abstract: Translocation, introduction, reintroduction, and assisted migrations are species conservation strategies that are 

attracting increasing attention, especially in the face of climate 

change. However, preventing the extinction of the suite of dependent species whose host species are threatened is seldom 

considered, and the effects on dependent species of moving 

threatened hosts are unclear. There is no published guidance 

on how to decide whether to move species, given this uncertainty. We examined the dependent-host system of 4 disparate 

taxonomic groups: insects on the feather-leaf banksia (Banksia 

brownii), montane banksia (B. montana), and Stirling Range 

beard heath (Leucopogon gnaphalioides); parasites of wild cats; 

mites and ticks on Duvaucel’s gecko (Hoplodactylus duvaucelii) 

and tuatara (Sphenodon punctatus); and internal coccidian parasites of Cirl Bunting (Emberiza cirlus) and Hihi (Notiomystis 

cincta). We used these case studies to demonstrate a simple 

process for use in species- and community-level assessments 

of efforts to conserve dependents with their hosts. The insects 

dependent on Stirling Range beard heath and parasites on tigers 

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 169

dominant species by locally rare species may require decades, 

and extinctions may occur when plant species cannot migrate 

fast enough to escape the consequences of climate change. 

Such lags may impair ecosystem services, such as carbon sequestration and clean water production. Thus, to assess global 

change, simulation of plant migration and local vegetation 

change by dynamic global vegetation models (DGVMs) is 

critical, yet fraught with challenges. Global vegetation models cannot simulate all species, necessitating their aggregation 

into plant functional types (PFTs). Yet most PFTs encompass 

the full spectrum of migration rates. Migration processes span 

scales of time and space far beyond what can be confidently 

simulated in DGVMs. Theories about climate change and 

migration are limited by inadequate data for key processes at 

short and long time scales and at small and large spatial scales. 

These theories must be enhanced to incorporate species-level 

migration and succession processes into a more comprehensive definition of PFTs.

Link: http://bio.huji.ac.il/ese/PDFs/Neilson_etalBioScience2005.pdf

477. Notaro M, Mauss A, Williams JW. 2012.

Projected vegetation changes for the American Southwest: combined dynamic modeling and bioclimatic-envelope approach. 

Ecological Applications 22:1365-1388.

Type: Journal 

Geographic Area: Southwestern USA

Keywords: bioclimatic envelope model, biodiversity, climate 

change, dynamic global vegetation model, Southwest United 

States 

Abstract: This study focuses on potential impacts of 21st 

century climate change on vegetation in the Southwest United 

States, based on debiased and interpolated climate projections 

from 17 global climate models used in the Fourth Assessment 

Report of the Intergovernmental Panel on Climate Change. 

Among these models a warming trend is universal, but projected changes in precipitation vary in sign and magnitude. 

Two independent methods are applied: a dynamic global vegetation model to assess changes in plant functional types and 

bioclimatic envelope modeling to assess changes in individual 

tree and shrub species and biodiversity. The former approach 

investigates broad responses of plant functional types to climate change, while considering competition, disturbances, and 

carbon fertilization, while the latter approach focuses on the 

response of individual plant species, and net biodiversity, to 

climate change. The dynamic model simulates a region-wide 

reduction in vegetation cover during the 21st century, with 

a partial replacement of evergreen trees with grasses in the 

mountains of Colorado and Utah, except at the highest elevations, where tree cover increases. Across southern Arizona, 

central New Mexico, and eastern Colorado, grass cover declines, in some cases abruptly. Due to the prevalent warming 

observed ~3 °C increase in minimum temperatures. Formerly 

low-elevation species expanded their ranges and high-elevation species contracted theirs, leading to changed community 

composition at mid- and high elevations. Elevational replacement among congeners changed because species’ responses 

were idiosyncratic. Though some high elevation species are 

threatened, protection of elevation gradients allows other species to respond via migration.

Link: http://www.nwf-wcr.org/PDFs-ClimateChangeFWP/

NRNRC-GW-FWP-Moritz-Paper.pdf

475. Mueller JM, Hellmann JJ. 2008. An assessment 

of invasion risk from assisted migration. Conservation Biology 

22:562-567.

Type: Journal 

Geographic Area: USA

Keywords: assisted migration, climate change, extinction 

risk, geographic distribution, invasive species, translocation

Abstract: To reduce the risk of extinction due to climate 

change, some ecologists have suggested human-aided translocation of species, or assisted migration (AM), to areas where 

climate is projected to become suitable. Such intentional 

movement, however, may create new invasive species if successful introductions grow out of control and cause ecologic or 

economic damage. We assessed this risk by surveying invasive 

species in the United States and categorizing invaders based on 

origin. Because AM will involve moving species on a regional 

scale within continents (i.e., range shifts), we used invasive 

species with an intracontinental origin as a proxy for species 

that would be moved through AM. We then determined whether intracontinental invasions were more prevalent or harmful 

than intercontinental invasions. Intracontinental invasions occurred far less frequently than invasions from other continents, 

but they were just as likely to have had severe effects. Fish and 

crustaceans pose a particularly high threat of intracontinental 

invasion. We conclude that the risk of AM to create novel invasive species is small, but assisted species that do become 

invasive could have large effects. Past experience with species 

reintroductions may help inform policy regarding AM.

Link: http://www.ncbi.nlm.nih.gov/pubmed/18577085

476. Neilson RP, Pitelka LF, Solomon AM, Nathan 

R, Midgley GF, Fragoso JMV, Lischke H, Thompson 

K. 2005. Forecasting regional to global plant migration in response to climate change. BioScience 55:749-759.

Type: Journal 

Geographic Area: Global

Keywords: climate change, dispersal, migration, long-distance dispersal, dynamic global vegetation models

Abstract: The rate of future climate change is likely to exceed 

the migration rates of most plant species. The replacement of 

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170 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Link: http://www.ualberta.ca/~ahamann/publications/pdfs/

ONeill_et_al_2008.pdf

479. O’Neill GA, Berger V, Carlson M, Ukrainetz 

NK. 2013. Assisted migration adaptation trial. British 

Columbia, Canada: BC Ministry of Forests, Lands and Natural 

Resource Operations. 2 p

Type: Government Document

Geographic Area: Western North America

Keywords: assisted migration, AMAT, Canada, tree management, forestry

Background: Seedlings are planted in the western USA, 

British Columbia (BC) and Yukon each year. Many climatologists predict that the climate could be 3–4 °C warmer when 

those trees are harvested 60-80 years after planting. These 

changes to climate will expose trees to increased stress and 

health risks, compromising the many goods and services we 

receive from our forests. Consequently, BC Forest Service 

researchers, with the assistance of the USDA Forest Service 

and industry, have initiated a large, long-term climate change 

research study—the Assisted Migration Adaptation Trial 

(AMAT)—to better understand tree species climate tolerances.

Link: https://www.for.gov.bc.ca/HRE/forgen/interior/AMAT.

htm

480. Parker KA, Seabrook-Davison M, Ewen JG. 

2008. Opportunities for nonnative ecological replacements in 

ecosystem restoration. Restoration Ecology 18:269-273.

Type: Journal 

Geographic Area: Australia

Keyword: Coturnix novaezelandiae, Coturnix ypsilophora, 

ecological analog, ecological restoration, translocation

Abstract: Translocations can take a variety of forms, and there 

is considerable debate as to what defines an acceptable translocation. This is particularly so if a proposal suggests moving 

a species beyond its natural range, which might be necessary 

for conservation purposes if habitat within the natural range is 

extensively modified. An extension of this approach is to use 

closely related ecological analogs to replace extinct species. 

This approach is controversial, and opportunities to do so will 

be rare, particularly for vertebrate species, but the use of ecological analogs is not without precedent, and ultimately will 

provide for more complete ecological restoration. We discuss 

the current use of ecological analogs to replace extinct species and conclude with a rare opportunity to replace the extinct 

New Zealand quail Coturnix novaezelandiae with the extant 

Australian brown quail Conturnix ypsilophora.

481. Parmesan C. 2006. Ecological and evolutionary responses to recent climate change. Annual Review of Ecology, 

Evolution, and Systematics 37:637-669.

trend among all 17 climate models, vegetation cover declines 

in the 21st century, with the greatest vegetation losses associated with models that project a drying trend. The inclusion of 

the carbon fertilization effect largely ameliorates the projected 

vegetation loss. Based on bioclimatic envelope modeling for 

the 21st century, the number of tree and shrub species that are 

expected to experience robust declines in range likely outweighs the number of species that are expected to expand in 

range. Dramatic shifts in plant species richness are projected, 

with declines in the high-elevation evergreen forests, increases 

in the eastern New Mexico prairies, and a northward shift of 

the Sonoran Desert biodiversity maximum. 

478. O’Neill GA, Hamann A, Wang T. 2008.

Accounting for population variation improves estimates of the 

impact of climate change on species’ growth and distribution.

Journal of Applied Ecology 45:1040-1049.

Type: Journal 

Geographic Area: Western Canada

Keywords: adaptation, assisted migration, ecological modelling, forest productivity, growth response function, Pinus contorta, population differentiation, seed transfer, transfer function

Abstract: Large differences exist in the predictions of plant 

responses to climate change among models that consider 

population variation and those that do not. Models that treat 

species as homogeneous entities typically predict net positive 

impacts of climate change on temperate forest productivity, 

while most studies that consider adaptive genetic variation 

within species conclude that the impacts of climate change on 

forest productivity will be negative. We present a modelling 

approach that predicts plant responses to climate change using 

both ecological and genetic variables. The model uses growth 

data from multi-site provenance trials together with climate 

data for provenance source locations and test sites to predict 

distribution and productivity of tree populations under climate 

change. We used an extensive lodgepole pine Pinus contorta 

provenance data set to illustrate the model. Spatially explicit 

predictions of the impacts of climate change on production

were developed and suggested that different populations of 

lodgepole pine will respond very differently to climate change. 

Large production losses will be seen in many areas, although 

modest production increases may occur in some areas by 2085. 

The model further projects a significant redistribution of the 

species’ potential habitat northwards and upwards in altitude 

over the next several decades. This study points to the need 

to consider population differences when modelling biotic responses to climate change, and suggests that climate change 

will render populations maladapted in many areas. The model 

also provides a key tool potentially to mitigate climate change 

impacts by identifying populations expected to be best adapted 

throughout the next generation of forests. Finally, the study 

highlights the value of wide-ranging long-term provenance 

tests in addressing key issues in ecology and climate change.

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 171

diagnostic fingerprint was found for 279 species. This suite of 

analyses generates ‘very high confidence’ (as laid down by the 

IPCC) that climate change is already affecting living systems. 

Link: http://www.geo.utexas.edu/courses/387H/Lectures/parmesan.pdf

483. Parmesan C, Ryrholm N, Stefanescu C, Hill 

JK, Thomas CD, Descimon H, Huntley B, Kaila L, 

Kullberg J, Tammaru T, Tennent WJ, Thomas JA, 

Warren M. 1999. Poleward shifts in geographical ranges 

of butterfly species associated with regional warming. Nature 

399:579-583.

Type: Journal 

Geographic Area: Europe

Compilers’ Keywords: climate change, range shifts, global 

temperature, migration

Abstract: Mean global temperatures have risen this century, 

and further warming is predicted to continue for the next 

50–100 years. Some migratory species can respond rapidly to 

yearly climate variation by altering the timing or destination of 

migration, but most wildlife is sedentary and so is incapable 

of such a rapid response. For these species, responses to the 

warming trend should be slower, reflected in poleward shifts 

of the range. Such changes in distribution would occur at the 

level of the population, stemming not from changes in the pattern of individuals’ movements, but from changes in the ratios 

of extinctions to colonizations at the northern and southern 

boundaries of the range. A northward range shift therefore 

occurs when there is net extinction at the southern boundary 

or net colonization at the northern boundary. However, previous evidence has been limited to a single species or to only a 

portion of the species’ range. Here we provide the first largescale evidence of poleward shifts in entire species’ ranges. In 

a sample of 35 non-migratory European butterflies, 63% have 

ranges that have shifted to the north by 35–240 km during this 

century, and only 3% have shifted to the south.

484. Parolo G, Rossi G. 2008. Upward migration of vascular plants following a climate warming trend in the Alps. Basic 

and Applied Ecology 9:100-107.

Type: Journal 

Geographic Area: Italy, Europe

Compilers’ Keywords: alpine-nival ecosystems, climate 

change, diaspora, historical records, long-term observation, 

monitoring, species richness, wind-dispersion, alpine plants

Summary: The aim of this study was to understand (1) whether 

warmer climatic conditions affected the vascular plant species 

composition, (2) the magnitude and rate of altitudinal changes 

in species distributions, and (3) whether an upward migration 

of alpine plants is connected to wind dispersal of diaspores. 

We compared historical records (1954–1958) with results from 

Type: Journal 

Geographic Area: Global

Keywords: aquatic, global warming, phenology, range shift, 

terrestrial, trophic asynchrony

Abstract: Ecological changes in the phenology and distribution of plants and animals are occurring in all well-studied marine, freshwater, and terrestrial groups. These observed changes 

are heavily biased in the directions predicted from global warming and have been linked to local or regional climate change 

through correlations between climate and biological variation, 

field and laboratory experiments, and physiological research. 

Range-restricted species, particularly polar and mountaintop 

species, show severe range contractions and have been the first 

groups in which entire species have gone extinct due to recent 

climate change. Tropical coral reefs and amphibians have been 

most negatively affected. Predator-prey and plant-insect interactions have been disrupted when interacting species have 

responded differently to warming. Evolutionary adaptations 

to warmer conditions have occurred in the interiors of species’ 

ranges, and resource use and dispersal have evolved rapidly at 

expanding range margins. Observed genetic shifts modulate local effects of climate change, but there is little evidence that 

they will mitigate negative effects at the species level.

Link: http://www.law.arizona.edu/depts/ele/AdaptationConference/PDFs/ParmesanAREES_Impacts2006.pdf

482. Parmesan C, Yohe G. 2003. A globally coherent 

fingerprint of climate change impacts across natural systems.

Nature 421:37-42.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: migration, biological trends, IPCC, 

range shifts, meta-analysis, plants, animals, phenology, 

modelling

Abstract: Casual attribution of recent biological trends to climate change is complicated because non-climatic influences 

dominate local, short-term biological changes. Any underlying 

signal from climate change is likely to be revealed by analyses 

that seek systematic trends across diverse species and geographic regions; however, debates within the Intergovernmental 

Panel on Climate Change (IPCC) reveal several definitions of 

a ‘systematic trend’. Here, we explore these differences, apply diverse analyses to more than 1,700 species, and show that 

recent biological trends match climate changes predictions. 

Global meta-analyses documented significant range shifts averaging 6.1 km per decade towards the poles (or metres per 

decade upward), and significant mean advancement of spring 

events by 2.3 days per decade. We define a diagnostics fingerprint of temporal and spatial ‘sign-switching’ responses 

uniquely predicted by twentieth century climate trends. Among 

appropriate long-term/large-scale/multi-species data sets, this 

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172 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Abstract: Climate change presents a potentially severe threat 

to biodiversity. Species will be required to disperse rapidly 

through fragmented landscapes in order to keep pace with the 

changing climate. An important challenge for conservation is 

therefore to manage landscapes so as to assist species in tracking 

the environmental conditions to which they are adapted. Here 

we develop a stochastic spatially explicit model to simulate 

plant dispersal across artificial fragmented landscapes. Based 

on certain assumptions as to the dispersal mechanism, we assess the impact that varying potential for rare long-distance 

dispersal (LDD) has on the ability to move over landscapes 

with differing spatial arrangements of suitable habitat clumped 

versus fragmented). Simulations demonstrate how the relative 

importance of landscape structure in determining migration 

ability may decrease as the potential for LDD increases. Thus, 

if LDD is the principal mechanism by which rapid large-scale 

migrations are achieved, strategically planned networks of 

protected habitat may have a limited impact on rates of largescale plant migrations. We relate our results to conventional 

principles for conservation planning and the geometric design 

of reserves, and demonstrate how reversal of these principles 

may maximise the potential for conservation under future 

climates. In particular, we caution against the justification of 

large-scale corridors on grounds of climate change since migration along corridors by standard dispersal mechanisms is 

unlikely to keep pace with projected change for many species. 

An improved understanding of the dispersal mechanisms by 

which species achieve rapid migrations, and the way that these 

processes are affected by patterns of landscape fragmentation, 

will be important to inform future conservation strategies.

Link: http://www.agci.org/dB/PDFs/05S3_RPearson_TDawson_conservation%20targets_0722.pdf

487. Phillips SJ, Anderson RP, Schapire RE. 2006.

Maximum entropy modeling of species geographic distributions. 

Ecological Modelling 190:231-259.

Type: Journal 

Geographic Area: Global

Keywords: maximum entropy, distribution, modeling, niche, 

range

Abstract: The availability of detailed environmental data, together with inexpensive and powerful computers, has fueled a 

rapid increase in predictive modeling of species environmental 

requirements and geographic distributions. For some species, 

detailed presence/absence occurrence data are available, allowing the use of a variety of standard statistical techniques. 

However, absence data are not available for most species. In 

this paper, we introduce the use of the maximum entropy method (Maxent) for modeling species geographic distributions 

with presence-only data. Maxent is a general-purpose machine 

learning method with a simple and precise mathematical formulation, and it has a number of aspects that make it well-suited 

recent plant surveys (2003–2005) from alpine to nival ecosystems in the Rhaetian Alps, N-Italy. The presence of all vascular 

plant species and their maximum altitude were recorded along 

a continuous altitudinal transect of 730 m. An increase in species richness from 153 to 166 species was observed. Moreover, 

52 species were recorded from altitudes 30–430 m higher than 

their 1950s limits, which corresponds to a median migration 

rate of 23.9 m/decade. In order to explain the observed migrations, the species wind-dispersal ability (diaspore weight and 

morphology) and the air temperature variation from 1926 to 

2003 were considered. Species with more pronounced altitudinal shifts possess lighter diaspores. The highest increase in 

species richness was found between 2,800 and 3,100 m a.s.l.; 

this appears to be related to an estimated shift of the permafrost 

limit by +240 m during the last 50 years. The mean air temperature in the region rose by +1.6 °C in summer and by +1.1 

°C in winter within this period. Climate warming is therefore 

considered as a primary cause of the observed upward migration of high mountain plants. Calculated altitudinal migration 

rates, however, varied remarkably among species. This would 

imply differential abilities of species to persist in an increasingly warmer climate. Species-specific conservation measures, 

including ex situ conservation, may therefore be required.

485. Pearson RG. 2006. Climate change and the migration capacity of species. Trends in Ecology and Evolution 

21:111-113.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: pollen, species and specificity, review, trees, migration rates

Abstract: In a recent paper, McLachlan et al. presented evidence that migration rates of two tree species at the end of the 

last glacial (c. 10-20 thousand years ago) were much slower 

than was previously thought. These results provide an important insight for climate-change impacts studies and suggest that 

the ability of species to track future climate change is limited. 

However, the detection of late-glacial refugia close to modern 

range limits also implies that some of our most catastrophic 

projections might be overstated.

Link: http://www.ncbi.nlm.nih.gov/pubmed/16701483

486. Pearson RG, Dawson TP. 2005. Long-distance 

plant dispersal and habitat fragmentation: identifying conservation targets for spatial landscape planning under climate change. 

Biological Conservation 123:389-401.

Type: Journal 

Geographic Area: Global

Keywords: long-distance dispersal, climate change, habitat 

fragmentation, spatially explicit model, systematic conservation planning

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dispersal into ESSP (115-3132 m). The findings indicate intensive mixing of the genes in this expanding population instead 

of founder effects and support the view that genetic diversity 

can be maintained in a long-lived species during rapid population expansion driven by climate warming.

Link: http://www.ncbi.nlm.nih.gov/pubmed/21199030

489. Potter KM, Hargrove WW. 2012. Determining 

suitable locations for seed transfer under climate change: a global quantitative method. New Forests 43:581-599.

Type: Journal 

Geographic Area: Southeastern USA

Keywords: restoration, conservation genetics, quantitative 

ecoregions, multivariate clustering, seed sources, human-assisted migration

Abstract: Changing climate conditions will complicate efforts to match seed sources with the environments to which 

they are best adapted. Tree species distributions may have to 

shift to match new environmental conditions, potentially requiring the establishment of some species entirely outside of 

their current distributions to thrive. Even within the portions 

of tree species ranges that remain generally suitable for the 

species, local populations may not be well-adapted to altered 

local conditions. To assist efforts to restore forests and to 

maximize forest productivity in the face of climate change, we 

developed a set of 30,000 quantitatively defined seed transfer 

“ecoregions” across the globe. Reflecting current and future 

conditions, these were created by combining global maps of 

potentially important environmental characteristics using a 

large-scale statistical clustering technique. This approach assigns every 4 km2 terrestrial raster cell into an ecoregion using 

nonhierarchical clustering of the cells in multivariate space 

based on 16 environmental variables. Two cells anywhere on 

the map with similar combinations of environmental characteristics are located near each other in this data space; cells are 

then classified into relatively homogeneous ecoregion clusters. 

Using two global circulation models and two emissions scenarios, we next mapped the predicted environmentally equivalent future locations of each ecoregion in 2050 and 2100. We 

further depicted areas of decreasing environmental similarity 

to given ecoregions, both in current time and under climate 

change. This approach could help minimize the risk that trees 

used for production, restoration, reforestation, and afforestation are maladapted to their planting site. 

Link: http://www.forestthreats.org/products/publications/Determining_suitable_locations_for_seed_transfer.pdf

490. Prunier J, Gerardi S, Laroche J, Beaulieu J, 

Bousquet J. 2012. Parallel and lineage-specific molecular 

adaptation to climate in boreal black spruce. Molecular Ecology 

21:4270-4286.

Type: Journal 

for species distribution modeling. In order to investigate the efficacy of the method, here we perform a continental-scale case 

study using two Neotropical mammals: a lowland species of 

sloth, Bradypus variegatus, and a small montane murid rodent, 

Microryzomys minutus. We compared Maxent predictions with 

those of a commonly used presence-only modeling method, the 

Genetic Algorithm for Rule-Set Prediction (GARP). We made 

predictions on 10 random subsets of the occurrence records for 

both species, and then used the remaining localities for testing. 

Both algorithms provided reasonable estimates of the species’ 

range, far superior to the shaded outline maps available in field 

guides. All models were significantly better than random in both 

binomial tests of omission and receiver operating characteristic 

(ROC) analyses. The area under the ROC curve (AUC) was almost always higher for Maxent, indicating better discrimination 

of suitable versus unsuitable areas for the species. The Maxent 

modeling approach can be used in its present form for many 

applications with presence-only datasets, and merits further research and development.

Link: http://perceval.bio.nau.edu/downloads/grail/climate_

seminar/section2/Phillips_Anderson_and_Schapire06.pdf

488. Pluess AR. 2011. Pursuing glacier retreat: genetic 

structure of a rapidly expanding Larix decidua population.

Molecular Ecology 20:473-485.

Type: Journal 

Geographic Area: Swiss Alps, Europe

Keywords: climate change, founder effects, glacier foreland, 

range expansion, spatial genetic structure, succession

Abstract: One of the greatest threats to the long-term viability of migrating plant species is the loss of genetic diversity 

due to founder effects. Populations can expand as a response 

to climate change, but it is uncertain if long lived plant species 

can maintain sufficient genetic diversity at the leading edge 

of migrating populations. This study uses an expanding Larix 

decidua population investigated along a chronosequence at 

landscape (350 ha) and local (0.8 ha) scales to test whether 

accelerated migration as a result of climate warming has the 

potential to intensify genetic erosion. Nine SSR markers revealed similar genetic diversity among eight sub-populations 

along the chronosequence (overall He = 0.73; SE = 0.04). Subpopulations were not genetically differentiated and all sampled 

individuals (N = 730) formed one major genetic cluster indicating homogenizing gene flow despite spatial genetic structure 

(SGS) up to 80 m. At the local scale, individuals at the leading 

edge [early successional sub-population (ESSP), N =140] and 

a sub-population at equilibrium [late successional sub-population (LSSP), N = 290] revealed high genetic diversity in 

largest-sized cohorts. SGS among juveniles occurred up to 30 

m in LSSP but there was no structure in ESSP. Accordingly, a 

maximum likelihood paternity assignment revealed local gene 

dispersal in LSSP (2-48 m) and intermediate-to-long distance 

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174 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

We used a spatially explicit forest ecosystem model, LANDISII, to simulate the interaction of climate change and forest management in northeastern Minnesota, USA. We assessed the relevance of restoration strategies and conservation targets based on 

the RNV in the context of future climate change. Three climate 

scenarios (no climate change, low emissions, and high emissions) were simulated with three forest management scenarios: 

no harvest, current management, and a restoration-based approach where harvest activity mimicked the frequency, severity, and size distribution of historic natural disturbance regimes. 

Under climate change there was a trend toward homogenization 

of forest conditions due to the widespread expansion of systems 

dominated by maple (Acer spp.). White spruce (Picea glauca), 

balsam fir (Abies balsamea), and paper birch (Betula papyrifera) 

were extirpated from the landscape irrespective of management 

activity; additional losses of black spruce (P. mariana), red pine 

(Pinus resinosa), and jack pine (P. banksiana) were projected 

in the high emissions scenario. In the restoration management 

scenario, retention and conversion to white pine (P. strobus) restricted maple expansion. But, widespread forest loss in the restoration scenario under high-emissions projections illustrates 

the potential pitfalls of implementing an RNV management approach in a system that is not compositionally similar to the historic reference condition. Given the uncertainty associated with 

climate change, ensuring a diversity of species and conditions 

within forested landscapes may be the most effective means of 

ensuring the future resistance of ecosystems to climate-induced 

declines in productivity.

Link: http://flux.aos.wisc.edu/~adesai/documents/macrosys_

papers-ankur/management/Ravenscroft_EcoApps_2010.pdf

492. Regan HM, Syphard AD, Franklin J, Swab 

RM, Markovchick L, Flint AL, Flint LE, Zedler PH. 

2012. Evaluation of assisted colonization strategies under 

global change for a rare, fire-dependent plant. Global Change 

Biology 18:936-947.

Type: Journal 

Geographic Area: California, USA

Keywords: assisted colonization, bioclimatic envelope, climate change, fire, population model

Abstract: As a clear consensus is emerging that habitat for 

many species will dramatically reduce or shift with climate 

change attention is turning to adaptation strategies to address 

these impacts. Assisted colonization is one such strategy that 

has been predominantly discussed in terms of the costs of 

introducing potential competitors into new communities and 

the benefits of reducing extinction risk. However, the success 

or failure of assisted colonization will depend on a range of 

population-level factors that have not yet been quantitatively 

evaluated—the quality of the recipient habitat, the number 

and life stages of translocated individuals, the establishment 

of translocated individuals in their new habitat and whether 

Geographic Area: North America

Keywords: adaptation, gene SNP, genetic lineages, natural 

selection, phylogeography, Picea mariana

Abstract: In response to selective pressure, adaptation may 

follow different genetic pathways throughout the natural range 

of a species due to historical differentiation in standing genetic variation. Using 41 populations of black spruce (Picea 

mariana), the objectives of this study were to identify adaptive 

genetic polymorphisms related to temperature and precipitation variation across the transcontinental range of the species, 

and to evaluate the potential influence of historical events on 

their geographic distribution. Population structure was first inferred using 50 control nuclear markers. Then, 47 candidate 

gene SNPs identified in previous genome scans were tested for 

relationship with climatic factors using an FST-based outlier 

method and regressions between allele frequencies and climatic variations. Two main intraspecific lineages related to glacial 

variance were detected at the transcontinental scale. Withinlineage analyses of allele frequencies allowed the identification of 23 candidate SNPs significantly related to precipitation 

and/or temperature variation, among which seven were common to both lineages, eight were specific to the eastern lineage 

and eight were specific to the western lineage. The implication of these candidate SNPs in adaptive processes was further 

supported by gene functional annotations. Multiple evidences 

indicated that the occurrence of lineage-specific adaptive SNPs 

was better explained by selection acting on historically differentiated gene pools rather than differential selection due to heterogeneity of interacting environmental factors and pleiotropic 

effects. Taken together, these findings suggest that standing 

genetic variation of potentially adaptive nature has been modified by historical events, hence affecting the outcome of recent 

selection and leading to different adaptive routes between intraspecific lineages.

Link: http://www.ncbi.nlm.nih.gov/pubmed/22805595

491. Ravenscroft C, Scheller RM, Mladenoff DJ, 

White MA. 2010. Forest restoration in a mixed-ownership 

landscape under climate change. Ecological Applications 

20:327-346.

Type: Journal 

Geographic Area: Minnesota, USA

Keywords: climate change, desired conditions, forest ecosystem model, LANDIS-II, forest management, historical variability, Minnesota, (USA), range of natural variability (RNV), 

restoration, species migration

Abstract: The extent to which current landscapes deviate from 

the historical range of natural variability (RNV) is a common 

means of defining and ranking regional conservation targets. 

However, climate induced shifts in forest composition may 

render obsolete restoration strategies and conservation targets 

based on historic climate conditions and disturbance regimes. 

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 175

for genetic differences among populations. Analyses also detected clines for the 19 P. pungens and 23 P. glauca populations. 

An assessment of the effects of global warming according to 

the IS92a scenario of two general circulation models demonstrated for the current century: (1) an increasingly favorable climate for P. pungens as its distribution moves upward 

in elevation throughout much of the Great Basin, Colorado 

Rockies, and mountain islands of the Southwest; (2) a widespread reduction in the areal extent of P. engelmannii in the 

inland Northwestern United States to the extent that Picea may 

become rare in the local flora; (3) extirpation of P. glauca from 

the Black Hills and Cypress Hills; and (4) a widespread redistribution of genotypes across the landscape as contemporary 

populations adjust genetically to change.

Link: http://www.treesearch.fs.fed.us/pubs/6952

494. Rehfeldt GE, Jaquish BC. 2010. Ecological impacts and management strategies for western larch in the face 

of climate-change. Mitigation and Adaptation Strategies for 

Global Change 15:283-306.

Type: Journal 

Geographic Area: Northwestern North America 

Keywords: bioclimate models, mapped genetic variation, 

seed zones, seed transfer guidelines, climate responses, climatic niche

Abstract: Approximately 185,000 forest inventory and ecological plots from both USA and Canada were used to predict 

the contemporary distribution of western larch (Larix occidentalis Nutt.) from climate variables. The random forests algorithm, using an 8 variable model, produced an overall error 

rate of about 2.9%, nearly all of which consisted of predicting 

presence at locations where the species was absent. Genetic 

variation among 143 populations within western larch’s natural distribution was predicted from multiple regression models 

using variables describing the climate of the seed source as 

predictors and response data from two separate genetic tests: 1) 

15-year height at a field site in British Columbia, Canada, and, 

2) two principal components of 8 variables describing growth, 

disease tolerance, and phenology of 6-year-old trees in a test in 

Idaho, USA. Presence and absence of the species and genetic 

variation within the species were projected into future climates 

provided by three General Circulation Models and two scenarios. Although the projections described pronounced impacts 

on the species and its populations, concurrence among the 

six projections pinpointed areas where the probability would 

be high that the future climate would be suitable for western 

larch. Concurrence among projections also was used to locate 

those sources of seed that should be best attuned genetically 

to future climates. The procedures outline a logical approach 

for developing management strategies for accommodating climate-change while taking into account the variability imposed 

by the differences among climatic estimates.

the recipient habitat is subject to ongoing threats all will play 

an important role in population persistence. In this article, we 

do not take one side or the other in the debate over whether 

assisted colonization is worthwhile. Rather, we focus on the 

likelihood that assisted colonization will promote population 

persistence in the face of climate-induced distribution changes 

and altered fire regimes for a rare endemic species. We link a 

population model with species distribution models to investigate expected changes in populations with climate change, the 

impact of altered fire regimes on population persistence and 

how much assisted colonization is necessary to minimize risk 

of decline in populations of Tecate cypress, a rare endemic tree 

in the California Floristic Province, a biodiversity hotspot. We 

show that assisted colonization may be a risk-minimizing adaptation strategy when there are large source populations that 

are declining dramatically due to habitat contractions, multiple 

nearby sites predicted to contain suitable habitat, minimal 

natural dispersal, high rates of establishment of translocated 

populations and the absence of nonclimatic threats such as 

altered disturbance regimes. However, when serious ongoing 

threats exist, assisted colonization is ineffective. 

493. Rehfeldt GE. 2004. Interspecific and intraspecific 

variation in Picea engelmannii and its congeneric cohorts: biosystematics, genecology, and climate change. Fort Collins (CO): 

USDA Forest Service, Rocky Mountain Research Station. 

General Technical Report RMRS-GTR-134. 22 p.

Type: Government Document

Geographic Area: Western North America

Keywords: common garden studies, quantitative traits, climatic clines, response to climate

Abstract: A series of common garden studies of 336 populations representing Picea engelmannii, P. pungens, P. glauca, P. 

mexicana, and P. chihuahuana provided as many as 13 growth 

and morphologic characters pertinent to biosystematics and 

genecology. Canonical discriminant analyses discretely segregated populations of P. pungens and P. chihuahuana while positioning P. engelmannii populations along a continuum anchored 

by Southwestern United States populations at one extreme and 

those classified as hybrids of P. engelmannii with P. glauca on 

the other. A population of P. mexicana was closely aligned with 

Southwest populations of P. engelmannii, while populations of 

P. glauca were intermixed with and peripheral to those identified as hybrid. While consistent with most taxonomic treatments of these taxa, the analyses nonetheless suggested that 

Southwestern United States populations should be considered 

as a variety of P. engelmannii that most likely should include 

P. mexicana. Genecological analyses detected ample genetic 

variation among the 295 populations in the P. engelmannii

complex. The analyses demonstrated that populations were 

distributed along clines driven primarily by the winter temperature regime of the provenance. For northern populations, 

summer temperatures also became a key factor in accounting 

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176 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

(2) developing population-specific response functions driven 

by predicted climate variables, (3) developing general transfer 

functions that predict performance from the climatic distances 

over which populations were transferred, and (4) interpreting the results in terms of niche breadth, effects of climate 

change on adaptedness of populations, and reforestation in a 

changing environment. Polynomial regression models used 

physiographic descriptors to predict seven climate variables 

from normalized records of 513 weather stations. Values of R2

ranged over 0.80–0.97 for thermal variables and 0.54–0.61 for 

precipitation variables. Validations with independent data from 

45 stations were strong and suggested that the models were 

generally free of bias within the limits of the original data. 

Response functions describing the height or survival of each 

population were developed from quadratic regressions using 

predicted climate variables for each test site. Mean annual temperature and mean temperature in the coldest month were the 

most effective variables for predicting population height, while 

the ratio of summer temperature to summer moisture was the 

best predictor of survival. Validation of the response functions 

with independent data from two additional test sites produced 

values of R2 between actual and predicted values that were as 

high as 0.93 for height and 0.73 for survival. The results demonstrated that natural populations have different climatic optima but tend to occupy suboptimal environments. Nevertheless, 

the general transfer functions showed that optimal growth and 

survival of the species as a whole is associated with the null 

transfer distance. These seemingly anomalous results suggest 

that the same processes thought to determine the distribution of 

species control the distribution of genotypes within species: (1) 

environmental selection to produce a broad fundamental niche, 

and (2) density-dependent selection to produce a relatively narrow realized niche within which most populations are relegate 

to suboptimal environments. Consequently, the steep geographic clines typical of P. contorta seem to be driven more by 

density-dependent selection than by environmental selection. 

Asymmetric gene flow from the center of distribution toward 

the periphery is viewed as a primary regulator that provides the 

fuel for both environmental and density dependent selection 

and thereby indirectly perpetuates suboptimality. The response 

functions predict that small changes in climate will greatly affect growth and survival of forest tree populations and, therefore, that maintaining contemporary forest productivities during global warming will require a wholesale redistribution of 

genotypes across the landscape. The response functions also 

provide the climatic bases to current reforestation guidelines 

and quantify the adjustments necessary for maintaining adaptedness in planted trees during periods of small (~1 ̊C) temporal 

temperature shifts.

497. Rehfeldt GE, Crookston NL, Warwell MV, 

Evans JS. 2006. Empirical analysis of plant-climate relationships for the western United States. International Journal of 

Plant Science 167:1123-1150.

Link: http://www.treesearch.fs.fed.us/pubs/35211

495. Rehfeldt GE, Wykoff WR, Ying CC. 2001.

Physiologic plasticity, evolution, and impacts of a changing climate on Pinus contorta. Climatic Change 50:355-376.

Type: Journal 

Geographic Area: Western North America

Compilers’ Keywords: Pinus contorta, lodgepole pine, forest productivity, adaptation

Abstract: Climate response functions for 125 Pinus contorta 

populations were updated to assess the impact of 16 climate 

change scenarios on forest productivity. Productivity was defined as the volume of wood expected per hectare at age 20 and 

was calculated as the product of predicted individual tree volumes, an initial stocking (1600 trees ha−1), and predicted survival. Impact was considered according to the transient effects 

of a changing climate governed by (1) physiological plasticity 

in the contemporary generation and (2) long-term evolutionary 

adjustments that provide adaptedness and optimize productivity in future generations. Direct short-term plastic responses 

were geographically complex and had repercussions throughout the species’ distribution even when temperature fluctuations were small (±1 °C) and changes in distribution were inconsequential. Evolutionary adjustments ameliorated negative 

short term impacts while enhancing the positive. Scenarios that 

encompassed predictions for global warming produced shortterm impacts that were negative in the south and positive in 

the north, but subsequent evolutionary adjustments projected 

substantial increases in productivity. The long-term adjustments may require only 1 to 3 generations in the north but 6 

to 12 generations in the south, thereby taking between 200 and 

1,200 years.

496. Rehfeldt GE, Ying CC, Spittlehouse DL, 

Hamilton DAJ. 1999. Genetic responses to climate in 

Pinus contorta: niche breadth, climate change, and reforestation.

Ecological Monographs 69:375-407.

Type: Journal 

Geographic Area: Western Canada

Keywords: climatic gradients, climate models, climate response functions of populations, fundamental niche vs. realized 

niche, genetic variation, global climate change, microevolution, niche breadth, Pinus contorta, population differentiation, 

reforestation in a changing environment, responses to a changing environment

Abstract: Fundamental plant–environment relationships were 

revealed by analyses of 20-yr height and survival of 118 populations representing two subspecies of Pinus contorta growing 

in common gardens at 60 environmentally disparate test sites 

in British Columbia. The approach involved (1) preparing 

models that described the general climate of British Columbia, 

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biomes. Errors of prediction from the statistical model averaged 

3.7%, but for individual biomes, ranged from 0% to 21.5%. In 

validating the ability of the model to identify climates without 

analogs, 78% of 1, 528 locations outside North America and 

81% of land area of the Caribbean Islands were predicted to 

have no analogs among the 46 biomes. Biome climates were 

projected into the future according to low and high greenhouse 

gas emission scenarios of three General Circulation Models for 

three periods, the decades surrounding 2030, 2060, and 2090. 

Prominent in the projections were (1) expansion of climates 

suitable for the tropical dry deciduous forests of Mexico, (2) 

expansion of climates typifying desert scrub biomes of western 

USA and northern Mexico, (3) stability of climates typifying 

the evergreen–deciduous forests of eastern USA, and (4) northward expansion of climates suited to temperate forests, Great 

Plains grasslands, and montane forests to the detriment of taiga 

and tundra climates. Maps indicating either poor agreement 

among projections or climates without contemporary analogs 

identify geographic areas where land management programs 

would be most equivocal. Concentrating efforts and resources 

where projections are more certain can assure land managers a 

greater likelihood of success.

Link: http://www.treesearch.fs.fed.us/pubs/40382

499. Rehfeldt GE, Tchebakova NM, Parfenova 

EI, Wykoff WR, Kuzmina NA, Milyutin LI. 2002.

Intraspecific responses to climate in Pinus sylvestris. Global 

Change Biology 8:912-929.

Type: Journal 

Geographic Area: North America, Europe, Asia

Keywords: climate-change impacts, climate response functions, microevolution, population adaptation, genetic responses to climate, predicting responses to change

Abstract: Five population-specific response functions were 

developed from quadratic models for 110 populations of Pinus 

sylvestris growing at 47 planting sites in Eurasia and North 

America. The functions predict 13 year height from climate: 

degree-days > 5 °C; mean annual temperature; degree-days 

< 0 °C; summer-winter temperature differential; and a moisture index, the ratio of degree-days > 5 °C to mean annual 

precipitation. Validation of the response functions with two 

sets of independent data produced for all functions statistically 

significant simple correlations with coefficients as high as 0.81 

between actual and predicted heights. The response functions 

described the widely different growth potentials typical of 

natural populations and demonstrated that these growth potentials have different climatic optima. Populations nonetheless 

tend to inhabit climates colder than their optima, with the disparity between the optimal and inhabited climates becoming 

greater as the climate becomes more severe. When driven by 

a global warming scenario of the Hadley Center, the functions 

described short-term physiologic and long-term evolutionary 

Type: Journal 

Geographic Area: Western USA

Keywords: bioclimatic models, Random Forests multipleregression tree, climatic distributions, climatic niche, response 

to climate change, global warming

Abstract: The Random Forests multiple-regression tree was 

used to model climate profiles of 25 biotic communities of the 

western United States and nine of their constituent species. 

Analyses of the communities were based on a gridded sample 

of ca. 140,000 points, while those for the species used presence-absence data from ca. 120,000 locations. Independent 

variables included 35 simple expressions of temperature and 

precipitation and their interactions. Classification errors for 

community models averaged 19%, but the errors were reduced 

by half when adjusted for misalignment between geographic 

data sets. Errors of omission for species specific models approached 0, while errors of commission were less than 9%. 

Mapped climate profiles of the species were in solid agreement 

with range maps. Climate variables of most importance for 

segregating the communities were those that generally differentiate maritime, continental, and monsoonal climates, while 

those of importance for predicting the occurrence of species 

varied among species but consistently implicated the periodicity of precipitation and temperature-precipitation interactions. 

Projections showed that unmitigated global warming should 

increase the abundance primarily of the montane forest and 

grassland community profiles at the expense largely of those 

of the subalpine, alpine, and tundra communities but also that 

of the arid woodlands. However, the climate of 47% of the 

future landscape may be extramural to contemporary community profiles. Effects projected on the spatial distribution of 

species-specific profiles were varied, but shifts in space and 

altitude would be extensive. Species-specific projections were 

not necessarily consistent with those of their communities.

Link: http://www.treesearch.fs.fed.us/pubs/25706

498. Rehfeldt GE, Crookston NL, Saenz-Romero 

C, Campbell EM. 2012. North American vegetation model 

for land-use planning in a changing climate: a solution to large 

classification problems. Ecological Applications 22:119-141.

Type: Journal

Geographic Area: North America

Keywords: climate change impacts, climate niche modeling, 

land management alternatives, Random Forests classification 

tree, vegetation models

Abstract: Data points intensively sampling 46 North American 

biomes were used to predict the geographic distribution of biomes from climate variables using the Random Forests classification tree. Techniques were incorporated to accommodate 

a large number of classes and to predict the future occurrence 

of climates beyond the contemporary climatic range of the 

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178 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

point of vulnerability within the lifecycle? Ecological Modelling 

249:50-58.

Type: Journal 

Geographic Area: Global

Keywords: spatially explicit, individual-based, dynamic 

model, dispersal, lifecycle, species distribution, functional 

traits, PPunCC

Abstract: Many organisms are threatened by global climate 

change, but as sessile organisms, plants are particularly vulnerable. Unable to individually move to areas with more suitable 

conditions as the climate changes, plants must instead rely on 

their seeds dispersing far and often in order to migrate fast 

enough to keep pace with the changing climate. The fragmentation of natural landscape by clearing for agricultural or urban 

development, or a species’ requirement for particular soil types 

or topography, makes this migration even more problematic. 

The likelihood of a plant species persisting will thus depend 

on factors such as the rate of climate change, the degree of 

landscape fragmentation, and also the plant species’ lifecycle, 

seed production, dispersal, and establishment. This paper presents a dynamic model, called PPunCC (Plant Persistence under Climate Change), that represents the important factors and 

processes likely to affect a plant species’ capacity to migrate 

across a landscape fast enough to avoid extinction. We use the 

model to show that creating large-scale corridors that increase 

the connectivity of fragmented landscapes through targeted 

restoration could help species migrate naturally and find suitable environments in new climates. We also examine the effect 

of varying the key point of vulnerability within its life cycle 

between juvenile establishment, adult mortality or seed production and conclude that this has a relatively minor effect on 

the prognosis for the plant and does not affect the predicted 

benefits of restoration.

502. Rice KJ, Emery NC. 2003. Managing microevolution: restoration in the face of global change. Frontiers in 

Ecology and the Environment 1:469-478.

Type: Journal 

Geographic Area: California, USA

Compilers’ Keywords: local adaptation, climate change, 

plant conservation genetics, review, vernal pools, blue oak 

woodlands

Abstract: Evidence is mounting that evolutionary change can 

occur rapidly and may be an important means by which species 

escape extinction in the face of global change. Consequently, 

biologists need to incorporate evolutionary thinking into management decisions in conservation and restoration ecology. 

Here, we review the genetic and demographic properties that 

influence the ability of populations to adapt to rapidly changing 

selective pressures. To illustrate how evolutionary thinking can 

influence conservation and restoration strategies, we compare 

effects that were geographically complex. The short-term impacts, enhance the positive, and in time, substantially increase 

productivity throughout most of the contemporary pine forests 

of Eurasia. Realizing the long-term gains will require redistribution of genotypes across the landscape, a process that should 

take up to 13 generations and therefore many years. 

Link: http://forest.akadem.ru/PDF/060322/tchebakova_9.pdf

500. Renton M, Shackelford N, Standish RJ. 2012.

Habitat restoration will help some functional plant types persist 

under climate change in fragmented landscapes. Global Change 

Biology 18:2057-2070.

Type: Journal 

Geographic Area: Australia

Keywords: assisted migration, corridors, dynamic model, 

functional traits, individual-based, intervention ecology, spatially-explicit, species distribution

Abstract: In the next century, global climate change is predicted to have large influences on species’ distributions. Much 

of the research in this area has focused on predicting the areas 

where conditions will be suitable for the species in future and 

thus the potential distribution of the species. However, it is 

equally important to predict the relative abilities of species to 

migrate into new suitable areas as conditions shift, while accounting for dynamic processes, such as dispersal, maturation, 

mortality, and reproduction, as well as landscape characteristics, such as level of habitat fragmentation and connectivity. In 

this study, we developed a spatially explicit individual-based 

model that addresses these factors. As a motivating case study, 

we based aspects of the model on southwest Australia, a global 

biodiversity hotspot, but stress that the results obtained are generalizable beyond this region. Using the model, we enhanced 

current understanding of climate change impacts by investigating how and to what extent the functional traits of plant species 

affect their ability to move with climate change across landscapes with various levels of fragmentation. We also tested the 

efficacy of strategic restoration, such as planting corridors to 

increase connectivity among fragments. We found that even 

if the landscape is fully intact, only an average of 34.2% of 

all simulated functional groups had a good change of successfully tracking climate change. However, our study highlights 

the power of strategic restoration as a tool for increasing species persistence. Corridors linking fragments increase species 

persistence rates by up to 24%. The lowest persistence rates 

were found for trees, a functional group with high dispersal but 

also long generation times. Our results indicate that for trees 

intervention techniques, such as assisted migration might be 

required to prevent species losses. 

501. Renton M, Childs S, Standish RJ, Shackelford 

N. 2013. Plant migration and persistence under climate 

change in fragmented landscapes: does it depend on the key 

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Geographic Area: California, USA

Compilers’ Keywords: review, plant taxa

Abstract: Based on our review of literature and survey of 

geneticists working on California taxa, we find genetic information lacking for most species in the Sierra Nevada. This 

situation is likely to remain in the future, with specific groups 

of taxa or occasional rare or high-interest species receiving 

specific study. Where we do have empirical information, we 

find few generalities emerging, except occasionally within 

closely related or ecologically similar taxa. Despite these difficulties in assessing genetic diversity, we direct attention to 

situations estimated to be most deserving of attention from a 

genetic standpoint.

Link: http://www.treesearch.fs.fed.us/pubs/6671

505. Saenz-Romero C, Rehfeldt GE, Crookston NL, 

Duval P, Beaulieu J. 2012. Spline models of contemporary, 2030, 2060 and 2090 climates for Michoacán state, Mexico.

Impacts on the vegetation. Revista Fitotecnica Mexicana 

35:333-345.

Type: Journal 

Geographic Area: Michoacán, Mexico

Index words: Pinus hartwegii, Pinus pseudostrobus, assisted 

migration, climate change, emission scenarios, Michoacán, 

thin plate smoothing splines

Summary: Climate data from 149 weather stations of 

Michoacán State, at Western México, were extracted from a 

spline climate model developed for México’s contemporary 

climate (1961–1990), and for climate projected for the decades 

centered in years 2030, 2060 and 2090. The model was constructed using outputs from three general circulation models 

(GCMs: Canadian, Hadley and Geophysical Fluid Dynamics) 

from two emission scenarios (A “pessimistic” and B “optimistic”). Mean annual temperature (MAT), mean annual precipitation (MAP), annual degree days > 5 °C (DD5), and annual 

aridity index (DD50.5/MAP) were mapped for Michoacán at an 

1 km2 scale, and means were estimated averaging all weather 

stations. The state average in GCMs and emission scenarios 

point out that mean annual temperature would increase 1.4 °C 

by year 2030, 2.2 °C by year 2060 and 3.6 °C by year 2090; 

whereas annual precipitation would decrease 5.6 % by year 

2030, 5.9 % by year 2060 and 7.8 % by year 2090. Climate 

models can be used for inferring plant-climate relationships 

and for developing programs to counteract global warming 

effects. Climate variables were estimated also at Pinus hartwegii and Pinus pseudostrobus growth locations, at Pico de 

Tancítaro in Central Western Michoacán and Nuevo San Juan 

Parangaricutiro (near Tancítaro), respectively. According to 

the annual aridity index values estimated for such locations, 

it is necessary to conduct assisted migration to match current 

genotypes to projected climates. This translates into an altitudinal shift of 400 to 450 m higher to match 2030 climates 

the potential of two California plant communities (vernal 

pools and blue oak woodlands) to evolve in response to global 

change. We then suggest ways in which restoration biologists 

can manipulate the genetic architecture of target populations 

to increase their ability to adapt to changing conditions. While 

there may not be any universal rules regarding the adaptive 

potential of species, an understanding of the various processes 

involved in microevolution will increase the short- and longterm success of conservation and restoration efforts.

Link: https://www.bio.purdue.edu/people/faculty/faculty_

files/publications/36121_1838168986.PDF

503. Richardson BA, Meyer SE. 2012. Paleoclimate effects and geographic barriers shape regional population genetic 

structure of blackbrush (Coleogyne ramosissima: Rosaceae).

Botany 90:293-299.

Type: Journal 

Geographic Area: Southwestern USA

Keywords: amplified fragment length polymorphisms, biogeography, Colorado Plateau, Mojave, postglacial colonization

Abstract: Coleogyne ramosissima Torr. (blackbrush) is a 

dominant xerophytic shrub species in the ecotone between 

the warm and cold deserts of interior western North America. 

Amplified fragment length polymorphisms (AFLPs) were 

used to survey genetic diversity and population genetic structure at 14 collection sites across the species range. Analysis 

revealed significant population differentiation (FST = 0.103, 

p < 0.0001) and reasonably high levels of genetic diversity 

(expected heterozygosity; HE = 0.26), a surprising result for 

a putative paleoendemic species. Model-based Bayesian clustering, principal coordinates analysis, and neighbor-joining 

analysis all produced support for the existence of two metapopulations, the first centered on the Mojave Desert and the 

second on the Colorado Plateau. These genetic data, coupled 

with information from Late Pleistocene and Holocene packrat 

(genus Neotoma Say and Ord, 1825) middens, illustrate a demographic history in which eastern and western distributions 

were disjunct during the Last Glacial Maximum and remained 

so through the Holocene, forming the present-day metapopulations in the Mojave Desert and Colorado Plateau. This strong 

regional genetic differentiation has implications for population 

persistence and migration in response to future climate change, 

as well as for shrubland restoration following anthropogenic 

disturbances such as annual grass invasion and wildfire.

Link: http://www.treesearch.fs.fed.us/pubs/40435

504. Rogers DL, Millar CI, Westfall RD. 1996.

Genetic diversity within species. Chapter 28. In: Erman DC, 

editor. Sierra Nevada Ecosystem Project: Final Report to 

Congress. Davis (CA): University of California, Davis. p 

759-838. 

Type: Book Section

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180 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Type: Journal 

Geographic Area: USA, Mexico

Compilers’ Keywords: climate change, GCM, Pinus 

chiapensis

Abstract: Spatial climate models were developed for 

México and its periphery (southern USA, Cuba, Belize and 

Guatemala) for monthly normals (1961–1990) of average, 

maximum and minimum temperature and precipitation using thin plate smoothing splines of ANUSPLIN software on 

ca. 3,800 observations. The fit of the model was generally 

good: the signal was considerably less than one-half of the 

number of observations, and reasonable standard errors for 

the surfaces would be less than 1 °C for temperature and 

10–15% for precipitation. Monthly normals were updated 

for three time periods according to three General Circulation 

Models and three emission scenarios. On average, mean annual temperature would increase 1.5 °C by year 2030, 2.3 °C 

by year 2060 and 3.7 °C by year 2090; annual precipitation 

would decrease -6.7% by year 2030, -9.0% by year 2060 

and -18.2% by year 2090. By converting monthly means 

into a series of variables relevant to biology (e.g., degreedays > 5 °C, aridity index), the models are directly suited 

for inferring plant–climate relationships and, therefore, in 

assessing impact of and developing programs for accommodating global warming. Programs are outlined for (a) assisting migration of four commercially important species of 

pine distributed in altitudinal sequence in Michoacán State 

(b) developing conservation programs in the floristically diverse Tehuacán Valley, and (c) perpetuating Pinus chiapensis, a threatened endemic. Climate surfaces, point or gridded 

climatic estimates and maps are available at http://forest.

moscowfsl.wsu.edu/climate/.

Link: http://www.treesearch.fs.fed.us/pubs/36311

508. Schwartz MW. 1992. Modelling effects of habitat fragmentation on the ability of trees to respond to climatic 

warming. Biodiversity and Conservation 2:51-61.

Type: Journal 

Geographic Area: Eastern North America

Keywords: tree migration, deforestation, Holocene, simulation model, global warming

Abstract: The ability of trees to migrate in response to climatic warming was simulated under various conditions of 

habitat availability. The model uses Holocene tree migration 

rates to approximate maximum migration rates in a forested 

landscape. Habitat availability and local population size was 

varied systematically under two dispersal and colonization 

models. These dispersal models varied in the likelihood 

of long-distance dispersal events. The first model used a 

negative exponential function that severely limited the probability of long-distance dispersal. The results of this model 

predicted by Canadian Model scenario A2, and 600 to 800 m 

to match 2060 climates.

Link: http://scielo.unam.mx/pdf/rfm/v35n4/v35n4a10.pdf

506. Sáenz-Romero C, Rehfeldt GE, Duval P, 

Lindig-Cisneros RA. 2012. Abies religiosa habitat 

prediction in climatic change scenarios and implications for 

monarch butterfly conservation in Mexico. Forest Ecology and 

Management 275:98-106.

Type: Journal 

Geographic Area: Michoacán, Mexico

Keywords: Danaus plexippus, suitable climatic habitat, 

Random Forests classification tree, assisted migration, climate 

change impacts, responses to climate 

Abstract: Abies religiosa (HBK) Schl. & Cham. (oyamel fir) 

is distributed in conifer-dominated mountain forests at high 

altitudes along the Trans-Mexican Volcanic Belt. This fir is the 

preferred host for overwintering monarch butterfly (Danaus 

plexippus) migratory populations which habitually congregate 

within a few stands now located inside a Monarch Butterfly 

Biosphere Reserve. Our objectives were to predict and map 

the climatic niche for A. religiosa for contemporary and future 

(2030, 2060 and 2090) climates, suggest management strategies to accommodate climate changes, and discuss implications for conservation of monarch butterfly overwintering 

sites in Mexico. A bioclimate model predicting the presence 

or absence of A. religiosa was developed by using the Random 

Forests classification tree on forest inventory data. The model 

used six predictor variables and was driven primarily by the 

mean temperature of the warmest month, an interaction between summer precipitation to and winter temperatures, and 

the ratio of summer to annual precipitation. Projecting the contemporary climate niche into future climates provided by three 

General Circulation Models and two scenarios suggested that 

the area occupied by the niche should diminish rapidly over the 

course of the century: a decrease of 69.2% by the decade surrounding 2030, 87.6% for that surrounding 2060, and 96.5% 

for 2090. We discuss assisted migration of A. religiosa upwards 

in altitude by 275 m so that populations of 2030 would occupy 

the same climates as today. The projections also show that by 

the end of the century, suitable habitat for the monarch butterfly may no longer occur inside the Biosphere Reserve. We 

therefore discuss management options and associated research 

programs necessary for assuring perpetuation of future butterfly habitat.

Link: http://www.treesearch.fs.fed.us/pubs/41323

507. Sáenz-Romero C, Rehfeldt GE, Crookston NL, 

Duval P, St-Amant R, Beaulieu J, Richardson BA. 

2009. Spline models of contemporary, 2030, 2060 and 2090 climates for Mexico and their use in understanding climate-change 

impacts on the vegetation. Climatic Change 102:595-623.

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 181

Geographic Area: Global

Keywords: adaptive evolution, additive genetic variation in 

fitness, geographic range, phenotypic plasticity, resurrection 

biology

Summary: Evolution proceeds unceasingly in all biological 

populations. It is clear that climate-driven evolution has molded 

plants in deep time and within extant populations. However, 

it is less certain whether adaptive evolution can proceed sufficiently rapidly to maintain the fitness and demographic stability 

of populations subjected to exceptionally rapid contemporary 

climate change. Here, we consider this question, drawing on 

current evidence on the rate of plant range shifts and the potential for an adaptive evolutionary response. We emphasize advances in understanding based on theoretical studies that model 

interacting evolutionary processes, and we provide an overview 

of quantitative genetic approaches that can parameterize these 

models to provide more meaningful predictions of the dynamic 

interplay between genetics, demography and evolution. We 

outline further research that can clarify both the adaptive potential of plant populations as climate continues to change and the 

role played by ongoing adaptation in their persistence.

Link: http://www.ncbi.nlm.nih.gov/pubmed/22816320

511. Smith SA, Beaulieu JM. 2009. Life history |influences rates of climatic niche evolution in flowering plants.

Proceedings of the Royal Society Biological Sciences 

276:4345-4352.

Type: Journal 

Geographic Area: Global

Keywords: life history, rates of evolution, phylogenetics, climatic niche

Abstract: Across angiosperms, variable rates of molecular 

substitution are linked with life-history attributes associated 

with woody and herbaceous growth forms. As the number of 

generations per unit time is correlated with molecular substitution rates, it is expected that rates of phenotypic evolution 

would also be influenced by differences in generation times. 

Here, we make the first broad-scale comparison of growthform-dependent rates of niche evolution. We examined the 

climatic niches of species on large time-calibrated phylogenies 

of five angiosperm clades and found that woody lineages have 

accumulated fewer changes per million years in climatic niche 

space than related herbaceous lineages. Also, climate space 

explored by woody lineages is consistently smaller than sister 

lineages composed mainly of herbaceous taxa. This pattern is 

probably linked to differences in the rate of climatic niche evolution. These results have implications for niche conservatism; 

in particular, the role of niche conservatism in the distribution 

of plant biodiversity. The consistent differences in the rate of 

climatic niche evolution also emphasize the need to incorporate models of phenotypic evolution that allow for rate heterogeneity when examining large datasets.

indicate that migration rate could decline an order of magnitude where the habitat availability is reduced from 80 to 20% 

of the matrix. The second model, using an inverse power 

function, carried a higher probability of long-distance dispersal events. The results from this model predict relatively 

small declines in migration rates when habitat availability is 

reduced to 50% of the simulation matrix. Below 50% habitat 

availability, mean migration rate was similar to the negative 

exponential model. These results predict a failure of many 

trees to respond to future climatic change through range 

expansion.

509. Schwartz MW. 1992. Potential effects of global 

climate change on the biodiversity of plants. The Forestry 

Chronicle 68:462-471.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: migration, trees, emissions, carbon 

dioxide

Abstract: Climatologists have observed a consistent increase in atmospheric CO2 over the past 30 years. It is 

predicted that CO2 levels could double the pre-industrial 

level of 280 ppm by the year 2100, perhaps much earlier. 

Climate models of doubled atmospheric CO2 predict that 

mean temperatures will increase between 1.5 and 4.5 °C 

globally; these temperature changes will be greater at high 

latitudes. Mid-continental regions will experience lower 

rainfall. Predictions of species northward range shifts in response to climate change vary from 100 km to over 500 km. 

Historical evidence of species range movements following 

the Pleistocene indicate that tree species typically migrated 

at rates of 10 km to 40 km per century. A simulation model 

that predicts the migration response of trees through modern fragmented landscapes predicts migration rates much 

lower than Pleistocene observations. Thus migration response is likely to lag far behind rates of climate change, 

potentially threatening narrowly distributed species whose 

predicted future ranges do not overlap with their current 

range. Insect pests and microbial pathogens should respond 

to climatic warming faster than long-lived trees. Predicted 

increased drought frequency may increase plant stress and 

thereby increase the frequency of insect outbreaks and disease. Predictions of species responses are complicated by 

direct effects of increased CO2 such as increased water-use 

efficiency. However, response to elevated CO2 varies among 

species. Thus, shifts in composition within plant communities are also likely, but are, as yet, unpredictable. 

510. Shaw RG, Etterson JR. 2012. Rapid climate change 

and the rate of adaptation: insight from experimental quantitative genetics. New Phytologist 195:752-765.

Type: Journal 

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182 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

efforts, while experimental common gardens should be established toward the south, to support active restoration approaches. This illustrates how ahead of time information on patterns 

of genetic variation can support restoration efforts for dryland 

tree species.

Link: http://www.researchgate.net/publication/233926307_

Genetic_Diversity_and_Structure_in_Austrocedrus_chilensis_Populations_Implications_for_Dryland_Forest_Restoration/file/79e4150d0aab77ac20.pdf

514. Spores S. 2009. Alaska yellow-cedar moves north. 

Juneau, AK: USDA Forest Service, Alaska Region. 3 p.

Type: Government Document

Geographic Area: Alaska

Compilers’ Keywords: facilitated migration, Tongass 

National Forest, yellow-cedar, assisted migration, forest 

management

Introduction: Climate change has become an increasingly 

prevalent topic in the news and in how the Forest Service 

will manage national forest lands for long term diversity and 

sustainability. The Forest Service has developed a strategic 

framework to guide management actions that address the challenges of climate change. One of the goals of this framework 

is “adaptation.” This means enhancing the capacity of forests 

to adapt to climate change in order to maintain ecosystem services. One of the methods that has been increasingly discussed 

is the idea of “facilitated migration,” or the moving of a species 

into an area where it doesn’t currently grow, but is anticipated 

to expand its range to that area in a warming climate. This kind 

of planting may be necessary because trees cannot migrate 

quickly enough keep up with the pace of anticipated climate 

change.

Link: http://www.fs.usda.gov/Internet/FSE_DOCUMENTS/

stelprdb5251356.pdf

515. St. Clair JB. 2013. Landscape variation in adaptation 

and implications for managing the future climates. In: Assisted 

Migration: A Primer for Reforestation and Restoration 

Decision Makers; Portland, OR. 136 p.

Type: Presentation

Geographic Area: Pacific Northwest USA

Compilers’ Keywords: assisted migration, forest management, Douglas-fir, genomics, provenance tests, bud burst

Abstract: Plant populations have evolved through natural 

selection to be genetically adapted to their local climates. 

Consequently, when climates change populations may become 

maladapted to the climate at their native location unless they 

can evolve through natural selection or migration at a pace 

that is fast enough to keep up with the change. Given the slow 

generation turnover of forest trees, along with other considerations, this seems unlikely. Assisted migration is an important 

Link: http://www.ncbi.nlm.nih.gov/pubmed/19776076

512. Smith JA, Trulock A. 2010. The decline of Florida 

torreya: an endemic conifer on the edge of extinction. Gainesville 

(FL): University of Florida. FOR276. 5 p.

Type: Report

Geographic Area: Southeastern USA

Compilers’ Keywords: species extinctions, endemic species

Overview: The Florida torreya (Torreya taxifolia) is a critically endangered conifer endemic to the Apalachicola River 

drainage of the Florida panhandle and adjacent southern 

Georgia. Florida torreya has experienced a significant dieoff and subsequent decline during the past century. Although 

the decline has been attributed to numerous abiotic and biotic 

causes, however, our findings are that the most likely reason 

for the decline is fungal disease. This fungal disease is likely 

caused by a Fusarium species and our suggestions for a new 

understanding of the actual cause of the decline are addressed 

in this fact sheet. Current efforts to assess the population and 

develop a recovery plan are discussed.

Link: http://edis.ifas.ufl.edu/pdffiles/FR/FR33800.pdf

513. Souto CP, Heinemann K, Kitzberger T, 

Newton AC, Premoli AC. 2011. Genetic diversity and 

structure in Austrocedrus chilensis populations: implications for 

dryland forest restoration. Restoration Ecology 20:568-575.

Type: Journal 

Geographic Area: Argentina, South America

Keywords: beyond-range restoration, fire, glaciations, passive restoration, Patagonia

Abstract: In South America, 94% of dry-temperate lands 

present some degree of environmental degradation, highlighting the need for ecological restoration. We analyzed geographic patterns of genetic variation in Austrocedrus chilensis, 

a dominant conifer of the steppe-forest ecotone in the eastern 

Andes, to examine its potential for restoration. We sampled 67 

locations in Argentina and estimated genetic parameters to determine the effects of historical factors affecting diversity, together with inbreeding and gene flow, using 12 allozyme loci. 

Genetic diversity decreased southwards in eastern populations, 

which are marginal for the range of the species and patchily 

distributed, while high genetic admixture was detected in continuous western populations, possibly reflecting postglacial 

migrations from northern and eastern sources. Higher inbreeding (FIS > 0.14) was recorded in northern compared with southern populations, attributed to the impact of recent bottlenecks 

resulting from anthropogenic fires. Gene flow was found to be 

moderate overall (FST = 0.12). The implications of these results 

for restoration on actions focusing on Austrocedrus were explored. Relatively small, inbred yet genetically diverse northern populations should be the subject of passive restoration 

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CHAPTER 3 • MIGRATION • RESEARCH

polymorphic loci (83.3%) and expected genetic diversity 

(0.427) in adult individuals were high; suggesting that all populations have genetic potential for conservation. The inbreeding within populations (ˆF = -0.011) and the total inbreeding 

(ˆf = 0.133) suggest population structure, since a high level of 

divergence among populations (θp = 0.143) was also detected. 

Significant values of spatial genetic structure were found inside 

the four populations. This study demonstrates that the realized 

gene flow among the remaining populations of O.catharinensis

is not sufficient to stop population divergence due to genetic 

drift and local selection, which threatens the future viability of 

this species.

517. Tchebakova NM, Rehfeldt GE, Parfenova EI. 

2010. From vegetation zones to climatypes: effects of climate 

warming on Siberian ecosystems. In: Osawa A, Zyryanova 

OA, Matsuura Y, Kajimoto T, Wein RW, editors. Permafrost 

Ecosystems: Siberian Larch Forests. 209. Netherlands: 

Springer. p 427-446. 

Type: Book Section

Geographic Area: Siberia, Asia

Compilers’ Keywords: climate change, range shifts, boreal 

forest, Pinus sylvestris, Larix sibirica

Introduction: Evidence for global warming over the past 

200 years is overwhelming, based on both direct weather observation and indirect physical and biological indicators such 

as retreating glaciers and snow/ice cover, increasing sea level, 

and longer growing seasons. Recent GCM projections of the 

Hadley Centre for Siberia show an increase in temperature of 

4 °C to 6 °C and an increase in precipitation of as much as 25% 

by 2100. These changes, moreover, could occur at a rate of 

0.1 °C to 0.4 °C per decade. The rapid rate of change coupled 

with the large absolute amount of change is expected to have 

profound effects on plants of the boreal forests at all hierarchical levels: from forest zones, to ecosystems, to species, to 

populations within species. Our goals are to estimate effects of 

a warming climate on Siberian vegetation, first, at the highest 

level of organization, and, second, the lowest. The first considers the effects of global warming as zonal vegetation shifts 

across the plains and plateaus of central Siberia, and the second 

considers intraspecific effects within the mountains of southern Siberia. For the second objective, we invoke Turesson’s 

concept of climatypes, the climatic ecotypes that comprise species, and illustrate intraspecific effects for Pinus sylvestris and 

Larix sibirica.

Link: http://www.treesearch.fs.fed.us/pubs/36298

518. Tchebakova NM, Parfenova EI, Soja AJ. 2011.

Climate change and climate-induced hot spots in forest shifts 

in central Siberia from observed data. Regional Environmental 

Change 11:817-827.

Type: Journal 

management option for maintaining adaptation, productivity 

and sustainability of forest ecosystems. Knowledge of landscape variation in adaption and adaptive traits is important for 

guiding decisions of population movement for purposes of assisted migration. Scientists study adaptation through genecology studies which consider correlations between population 

variation in traits and the environments of source locations 

where they evolved, or through reciprocal transplant studies 

which compare responses of different populations from a range 

of source environments grown in the same or similar range 

of environments. Results from genecology and reciprocal 

transplant studies of Douglas-fir are presented. Management 

options for responding to climate change are considered. One 

option is to do nothing, and it is important to consider the consequences of continuing to use current see zones and population movement guidelines. Silvicultural measures such as 

density management and fuels treatment may be an important 

option for maintaining resiliency and resisting change, but may 

just delay the inevitable as aging stands become increasingly 

maladapted. Managers may increase the likelihood of gene 

flow and migration by avoiding fragmentation and maintaining corridors for gene flow. But that may not be sufficient to 

keep pace with climate change. Assisted migration may be one 

of our most powerful tools for responding to climate change. 

Movement need not be geographically large at this point. The 

most critical phase of stand development is most likely to be 

the seedling and sampling stage; thus, movements should focus 

on the amount of climate change in the next couple of decades, 

generally, less than 2 °C. Movements of this magnitude are 

within current practices of transfers within current seed zones. 

Thus, risk is likely to be small, and managers should pay attention that movements are in the direction of warmer sources 

to cooler planting sites, and not both directions as is current 

practice. To facilitate assisted migration, seed zones and seed 

movement guidelines should be based on climate rather than 

geography, and seed collections should be bulked over a narrower climatic range than bulking using current seed zones. 

Link: http://www.rngr.net/resources/assisted-migration

516. Tarazi R, Mantovani A, dos Reis MS. 2009.

Fine-scale spatial genetic structure and allozymic diversity in 

natural populations of Ocotea catharinensis Mez. (Lauraceae). 

Conservation Genetics 11:965-976.

Type: Journal 

Geographic Area: Brazil, South America

Keywords: Atlantic rain forest, Lauraceae, endangered species, genetic diversity, spatial autocorrelation

Abstract: In order to establish a strategy for conservation, 

the distribution of genetic diversity in four natural populations of Ocotea catharinensis in the Brazilian Atlantic rain 

forest was investigated using 18 allozyme loci. Estimates of 

the average number of alleles per loci (2.2), percentage of 
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CHAPTER 3 • MIGRATION • RESEARCH

adapted to projected future climates than indigenous species. 

Here, we tested juvenile plants of 6 provenances of P. nigra for 

their response to different climate change scenarios (42-days 

drought and warming by 1.6K) in a full-factorial common-garden experiment in Bayreuth, Germany. In the second year only 

the warming treatment was imposed. Height, shoot mortality 

and needle phenology were determined for the two consecutive years. Provenances differed in absolute growth (from 6.0 

to 7.4 in 2009 and from 4.4 to 5.9 cm in 2010) and survival 

rates (from 78.6% to 97.6%), but not in terms of shoot quantity and, surprisingly, sensitivity to drought and warming. The 

drought treatment showed a delayed impact on height growth, 

as a significant growth reduction was detected for the second 

year (-2.6 cm), but not for the first year, when the actual treatment took places. Drought decreases survival rates by 20.6%. 

The drought treatment had no effect on needle phenology. 

Warming did not impact growth performance (height, shoot 

quantity). However, warming in combination with the drought 

treatment decreased the survival rate (-13%). Plants exposed 

to the warming treatment showed an earlier onset of needle 

development (-10.6 days). Our results imply that no significant 

local adaptation of growth and survival to drought and warming occurs in P. nigra. Performance and growth response in face 

of warming and drought cannot be predicted by the climate at 

the places of origin. We therefore recommend that an introduction of P. nigra to regions outside its natural distribution range 

should not aim at introducing a single best-adapted provenance 

but at establishing populations with a high genetic diversity, 

e.g. by promoting natural regeneration in native stands or by 

intermixing with different genotypes in order to maintain a 

high adaptive capacity to climate change. 

Link: http://www.researchgate.net/profile/Carl_Beierkuhnlein/publication/224758416_Uniform_drought_and_warming_responses_in_Pinus_nigra_provenances_despite_specific_overall_performances/links/0046351a5b0e193e7d000000.

pdf

520. Thorpe AS, Kaye TN. 2011. Conservation and reintroduction of the endangered Willamette daisy. Native Plants 

Journal 12:289-300.

Type: Journal 

Geographic Area: Oregon, USA

Keywords: reintroduction, Willamette Valley, prairie, endangered species, allee effect, Erigeron decumbens, Asteraceae

Abstract: Our research suggests that habitat fragmentation 

and reduced population size in Willamette daisy has led to loss 

of fitness in some populations and to population differentiation. In outcrossing species, habitat fragmentation and reduced 

population size can lead to increased genetic drift, local adaptation, and (or) inbreeding. We explored some of these issues 

for the endangered forb Willamette daisy (Erigeron decumbens Nutt. [Asteraceae]). This species is currently restricted 

Geographic Area: Siberia, Asia

Keywords: instrumental climate record, IPCC climate change 

projections, forest shift, Siberia 

Abstract: Regional Siberian studies have already registered 

climate warming over the last several decades. We evaluated 

ongoing climate change in central Siberia between 1991 and 

2010 and a baseline period, 1961–1990, and between 1991 and 

2010 and Hadley 2020 climate change projections, represented 

by the moderate B1 and severe A2 scenarios. Our analysis 

showed that winters are already 2–3 °C warmer in the north 

and 1–2 °C warmer in the south by 2010. Summer temperatures increased by 1 °C in the north and by 1–2 °C in the south. 

Change in precipitation is more complicated, increasing on 

average 10% in middle latitudes and decreasing 10–20% in the 

south, promoting local drying in already dry landscapes. Hot 

spots of possible forest shifts are modeled using our Siberian 

bioclimatic vegetation model and mountain vegetation model 

with respect to climate anomalies observed pre-2010 and predicted 2020 Hadley scenarios. Forests are predicted to shift 

northwards along the central Siberian Plateau and upslope in 

both the northern and southern mountains. South of the central 

Siberian Plateau, steppe advancement is predicted that was 

previously non-existent north of 56° N latitude. South of 56° 

N, steppe expansion is predicted in the dry environments of 

Khakasiya and Tyva. In the southern mountains, it is predicted 

that the lower tree line will migrate upslope due to increased 

dryness in the intermontane Tyvan basins. The hot spots of 

vegetation change that are predicted by our models are confirmed by regional literature data.

519. Thiel D, Nagy L, Beierkuhnlein C, Huber G, 

Jentsch A, Konnert M, Kreyling J. 2012. Uniform 

drought and warming responses in Pinus nigra provenances 

despite specific overall performances. Forest Ecology and 

Management 270:200-208.

Type: Journal 

Geographic Area: Germany, Europe

Keywords: European black pine, local adaptation, withinspecies diversity, provenance trial, plant-climate interactions, 

lag effect

Abstract: Climate extremes are expected to increase in frequency and magnitude as a consequence of global warming, 

threating the functioning, services and goods of forest ecosystems. The introduction of species from drier and warmer climates is one option that is discussed to adapt forest ecosystems 

to these adverse effects of climate change. The (sub)-Mediterranean Pinus nigra is a potential candidate for such assisted 

migration, especially for dry sites in Central Europe. The high 

genetic diversity within this species and thus the potential differences in adaptive capacity, however, makes it necessary to 

assess the response of P. nigra provenances to climatic extremes 

and identify suitable populations or ecotypes which are better 
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 185

patterns of projected climate change. This study highlights 

general patterns about the relationships between sensitivity 

of species to climate change and their ecological properties. 

There is a strong convergence between simple inferences 

based on ecological characteristics of species and projections 

by bioclimatic ‘envelope’ models, confirming macroecological 

assumptions about species sensitivity based on niche properties. These patterns appear to be most strongly driven by the 

exposure of species to climate change, with additional effects 

of species niche characteristics. We conclude that simple species niche properties are powerful indicators of species’ sensitivity to climate change.

Link: http://www.researchgate.net/publication/228851892_

Niche_properties_and_geographical_extent_as_predictors_of_species_sensitivity_to_climate_change/

file/79e4150c60275ab41b.pdf

522. Thuiller W, Albert C, Araújo MB, Berry 

PM, Cabeza M, Guisan A, Hickler T, Midgley GF, 

Paterson J, Schurr FM, Sykes MT, Zimmermann 

NE. 2008. Predicting global change impacts on plant species’ 

distributions: future challenges. Perspectives in Plant Ecology, 

Evolution and Systematics 9:137-152.

Type: Journal 

Geographic Area: Global

Keywords: species distribution modeling, habitat models, 

process-based models, global change, conservation planning

Abstract: Given the rate of projected environmental change 

for the 21st century, urgent adaptation and mitigation measures are required to slow down the on-going erosion of 

biodiversity. Even though increasing evidence shows that 

recent human-induced environmental changes have already 

triggered species’ range shifts, changes in phenology and 

species’ extinctions, accurate projections of species’ responses to future environmental changes are more difficult 

to ascertain. This is problematic, since there is a growing 

awareness of the need to adopt proactive conservation planning measures using forecasts of species’ responses to future 

environmental changes. There is a substantial body of literature describing and assessing the impacts of various scenarios of climate and land-use change on species’ distributions. 

Model predictions include a wide range of assumptions and 

limitations that are widely acknowledged but compromise 

their use for developing reliable adaptation and mitigation 

strategies for biodiversity. Indeed, amongst the most used 

models, few, if any, explicitly deal with migration processes, 

the dynamics of population at the ‘‘trailing edge’’ of shifting populations, species’ interactions and the interaction between the effects of climate and land-use. In this review, we 

propose two main avenues to progress the understanding and 

prediction of the different processes occurring on the leading and trailing edge of the species’ distribution in response 

to approximately 40 sites in the Willamette Valley, Oregon; 

more than half of the known populations have fewer than 100 

individuals. We found a positive relationship between seed 

viability (as measured by the percentage of filled seeds) and 

population size. In populations with 20 or fewer individuals, 

seed viability dropped to less than 2.5%. In modified reciprocal transplant and common-garden experiments, we found 

that although there were some differences in survival, growth, 

and reproduction in transplants from 2 source populations, the 

effect of source varied depending on response variable, year 

of planting, year of monitoring, and habitat. Conservation of 

Willamette daisy will benefit from increasing the size and genetic diversity of existing populations and from reintroducing 

genetically diverse populations within the historic range of the 

species. Knowledge of genetic diversity of populations can be 

critical for making appropriate management decisions for rare 

species, including determining if population augmentation is 

necessary and, if so, from which source populations.

Link: http://appliedeco.org/reports/Kaye%202008%20Reintroduction%20Native%20Plants%20Journal.pdf

521. Thuiller W, Lavorel S, Araujo MB. 2005. Niche 

properties and geographical extent as predictors of species sensitivity to climate change. Global Ecology and Biogeography 

14:347-357.

Type: Journal 

Geographic Area: Europe, Global

Keywords: bioclimatic ‘envelope’ modelling, climate change, 

future potential habitats, niche breadth, niche position, range 

size

Abstract: Bioclimatic envelope models are often used to 

make projections of species’ potential responses to climate 

change. It can be hypothesized that species with different 

kinds of distributions in environmental niche and geographical space may respond differently to changes in climate. Here, 

we compare projections of shifts in species ranges with simple 

descriptors of species niche (position and breadth) and geographical (range size) distributions. The future distribution for 

1,200 European plant species were predicted by niche-based 

models using seven climate variables known to have an important role in limiting plant species distributions. Ecological 

niche properties were estimated using a multivariate analysis. 

Species range changes were then related to species niche properties using generalized linear models. Generally, percentage 

of remaining suitable habitat in the future increased linearly 

with niche position and breadth. Increases in potential suitable habitat were associated with greater range size, and had 

a hump-shaped relationship with niche position on temperature gradient. By relating species chorotypes to percentage of 

remaining or gained habitat, we highlighted biogeographical 

patterns of species sensitivity to climate change. These were 

clearly related to the degree of exposure according to regional 

CHAPTER 3 • MIGRATION • RESEARCH
186 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

524. Trivedi MR, Morecroft MD, Berry PM, 

Dawson TP. 2008. Potential effects of climate change on 

plant communities in three montane nature reserves in Scotland, 

UK. Biological Conservation 141:1665-1675.

Type: Journal

Geographic Area: Scotland, United Kingdom

Keywords: Arctic-alpine plants, classification tree, Grampian 

highlands, mountains, Natura 2000, special area of conservation, species distribution models, topography

Abstract: Mountain ecosystems are often identified as being particularly sensitive to climate change, however this has 

rarely been investigated at the scale of individual mountain 

ranges using local relationships between plants and climate. 

This study uses fine resolution data to assess the potential 

changes to internationally important Arctic-alpine plant 

communities in three national nature reserves in the Scottish 

Highlands, United Kingdom. Distribution models were created for 31 species, representing a range of community types. 

A relationship between distribution and temperature was 

found for all species. These models were aggregated to explore potential future changes to each community under two 

Intergovernmental Panel on Climate Change warming scenarios for the 2080s. The results indicate that Arctic-alpine 

communities in these reserves could undergo substantial species turnover, even under the lower climate change scenario. 

For example, Racomitrium-Carex mossheath, a distinctive 

community type of the British uplands, could lose suitable 

climate space as other communities spread uphill. These 

findings highlight the need to maintain these communities 

in an optimal condition in which they can be most resilient 

to such change, to monitor them for signals of change and to 

develop more flexible conservation policies which account 

for future changes in mountain protected areas.

Link: https://edit.ethz.ch/ecology/education/BE_documents/

Trivedi_et_Al_2008_plant_community_shift_climate_change.

pdf

525. Ukrainetz NK, O’Neill GA, Jaquish B. 2011.

Comparison of fixed and focal point seed transfer systems 

for reforestation and assisted migration: a case study for interior spruce in British Columbia. Canadian Journal of Forest 

Research 41:1452-1464.

Type: Journal 

Geographic Area: Western Canada

Compilers’ Keywords: assisted colonization, assisted migration, maladaptation, Picea engelmannii, Picea glauca

Abstract: In forestry, science-based seed transfer systems, 

the foundation of effective reforestation programs, will likely be used in some form to mitigate the negative effects of 

climate change. In this study, we developed fixed and focal 

point seed transfer systems for interior spruce (Picea glauca

to any global change phenomena. Deliberately focusing on 

plant species, we first explore the different ways to incorporate species’ migration in the existing modelling approaches, 

given data and knowledge limitations and the dual effects 

of climate and land-use factors. Secondly, we explore the 

mechanisms and processes happening at the trailing edge of 

a shifting species’ distribution and how to implement them 

into a modelling approach. We finally conclude this review 

with clear guidelines on how such modelling improvements 

will benefit conservation strategies in a changing world.

Link: http://biology.mcgill.ca/grad/cecile/PDFs/3_

Thuiller2008.pdf

523. Tingley MW, Monahan WB, Beissinger SR, 

Moritz C. 2009. Birds track their Grinnellian niche through 

a century of climate change. Proceedings of the National 

Academy of Sciences USA 106 Supplement 2:19637-19643.

Type: Journal 

Geographic Area: California, USA

Keywords: climatic niche, geographic range, elevational gradient, occupancy dynamics

Abstract: In the face of environmental change, species can 

evolve new physiological tolerances to cope with altered 

climatic conditions or move spatially to maintain existing physiological associations with particular climates that 

define each species’ climatic niche. When environmental 

change occurs over short temporal and large spatial scales, 

agile species are expected to move geographically by tracking their climatic niches through time. Here, we test for evidence of niche tracking in bird species of the Sierra Nevada 

Mountains of California, focusing on 53 species resurveyed 

nearly a century apart at 82 sites on four elevational transects. Changes in climate and bird distributions resulted 

in focal species shifting their average climatological range 

over time. By comparing the directions of these shifts relative to the centroids of species’ range-wide climatic niches, 

we found that 48 species (90.6%) tracked their climatic 

niche. Analysis of niche sensitivity on an independent set of 

occurrence data significantly predicted the temperature and 

precipitation gradients tracked by species. Furthermore, in 

50 species (94.3%), site-specific occupancy models showed 

that the position of each site relative to the climatic niche centroid explained colonization and extinction probabilities better than a null model with constant probabilities. Combined, 

our results indicate that the factors limiting a bird species’ 

range in the Sierra Nevada in the early 20th century also 

tended to drive changes in distribution over time, suggesting that climatic models derived from niche theory might 

be used successfully to forecast where and how to conserve 

species in the face of climate change.

Link: http://www.ncbi.nlm.nih.gov/pubmed/19805037

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 187

Type: Journal 

Geographic Area: Venezuelan Guayana, South America 

Keywords: habitat loss, fragmentation, extinction, refugia, 

microrefugia, conservation, Venezuelan Guayana, neotropics

Abstract: Current global average temperatures are 2 °C 

cooler than during the last interglacial period. The expected 

increase in temperature during the 21st century will be most 

stressful for cold-adapted and stenothermic mountain species, 

forcing them to migrate upwards, and eventually to concentrate into either large areas with favourable climates (refugia) 

or small stands under locally favourable microclimates (microrefugia). We investigate potential refugia in the neotropical 

Guayana Highlands mountain biome (Pantepui), consisting 

of ~50 isolated table mountains (tepuis), to develop strategies 

for conserving biodiversity during future global warming. We 

predict the amount of loss of altitudinal habitats of endemic 

vascular flora of 26 tepuis and evaluate potential threats to 

these taxa with respect to species extinction, habitat loss, habitat connectivity and the degree of isolation. We compare past, 

present, and future Pantepui landscape configurations through 

fragmentation analysis and identify potential in situ refugia. 

Spatial analysis forecasts more species isolation and declining 

biodiversity at the end of this century relative to current and 

past levels. Habitats are predicted to experience > 80% loss, 

with the disappearance of 38 habitat patches. One large patch 

(Chimanta massif) accounts for 46% of the predicted remaining habitat. This patch can be considered a potential refugium 

for future vascular flora, as it is predicted to contain some 

present-day resistant species from lower altitudinal levels and 

other species eventually persisting in microrefugia. The easternmost Pantepui district, containing the Chimanta massif and 

other tepuis, seems to be the most suitable for the application 

of in situ conservation strategies.

528. Wagner M, Heinrich W, Jetschke G. 2006.

Seed bank assembly in an unmanaged ruderal grassland recovering from long-term exposure to  industrial emissions. Acta 

Oecologica 30:342-352.

Type: Journal 

Geographic Area: Germany, Europe

Keywords: derelict industrial ecosystem, grassland, 

Puccinellia distans, regeneration, secondary succession, seed 

bank composition, similarity patterns

Abstract: The main aim of this 3-year study was to relate 

the temporal patterns in seed bank composition of a ruderal 

grassland previously subjected to industrial pollution with 

the successional patterns in the above ground vegetation. 

In particular, we tested whether the observed changes conformed to a paradigm of declining seed numbers and diversity and decreasing similarity between seed bank and vegetation, as previously formulated for secondary successions. 

We found that seed numbers and the number of species per 

(Moench) Voss, Picea engelmannii Parry ex Engelm., and 

their hybrids) in British Columbia, Canada, and compared 

the effectiveness of both systems. Growth, phenology, and 

physiology traits were measured for 112 populations, and 

population means were transformed to principal components 

that were modeled using climate variables and multiple regression analysis. Compared with the fixed seed zone system, the focal point system had a greater area of seed use 

for a given risk of maladaptation. The relationship between 

growth and adaptive distance (i.e., adaptive similarity between test populations and populations local to test sites) 

was used to calculate critical seed transfer distances for focal 

point seed zones, which were defined according to expected 

forfeiture of growth. Changes in climate observed over the 

past 100 years and predicted in the next one third of a rotation were used to calculate appropriate assisted migration 

distances and develop methods for incorporating assisted 

migration into a focal point seed transfer system.

526. Van der Veken S, Hermy M, Vellend M, 

Knapen A, Verheyen K. 2008. Garden plants get a 

head start on climate change. Frontiers in Ecology and the 

Environment 6:212-216.

Type: Journal 

Geographic Area: Europe

Compilers’ Keywords: assisted migration, horticulture, 

commercial nurseries

Abstract: Conservation biologists are concerned that climate 

change will cause widespread extinctions because limited capacity for migration could compromise species’ ability to adjust 

to geographic shifts in habitat condition. However, commercial 

plant nurseries may provide a head start for northward range 

shifts among some plant species. To investigate this possibility, 

we compared the natural ranges of 357 native European plant 

species with their commercial ranges, based on 246 plant nurseries throughout Europe. In 73% of native species, commercial 

northern range limits exceeded natural northern range limits, 

with a mean difference of ~ 1000 km. With migration rates of 

~ 0.1–5 km per year required for geographic ranges to track 

climate change over the next century, we expect nurseries and 

gardens to provide a substantial head start on such migration 

for many native plants. While conservation biologists actively 

debate whether we should intentionally provide “assisted migration,” it is clear that we have already done so for a large 

number of species.

Link: http://www3.botany.ubc.ca/vellend/Van_der_Veken_

Frontiers2008.pdf

527. Vegas-Vilarrúbia T, Nogué S, Rull V. 2012.

Global warming, habitat shifts and potential refugia for biodiversity conservation in the neotropical Guayana Highlands.

Biological Conservation 152:159-168.

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188 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

and not as much due to geographic and climatic variables, 

metabolic profiles appeared best matched to the stress levels 

within local environments. It is unknown if differences in secondary metabolite levels are the result of genetic adaptation or 

phenotypic plasticity, but results from this and other studies 

suggest that both are important. These results are interpreted 

in light of ongoing efforts to assist in the migration of certain 

populations of P. c. latifolia northward in an effort to counter 

predicted effects of climate change.

Link: http://www.ncbi.nlm.nih.gov/pubmed/21537900

530. Wang T. 2012. Projecting future distributions of 

ecosystem climate niches in British Columbia. Journal of 

Ecosystems and Management 13:1-3.

Type: Journal 

Geographic Area: Western Canada

Compilers’ Keywords: climate change, random forest, bioclimate envelope modeling, IPCC, model projections, tree distribution, range shift, migration

Introduction: With accumulating evidence showing the 

ecological impacts of global climate change, scientists, 

land managers, and policymakers in British Columbia have 

become increasingly concerned about its impact on local 

ecosystems. One of the major concerns is the mismatch between the climate that an ecosystem is adapted to and the 

climate that the ecosystem will experience in the future. If 

such a mismatch occurs, the health and productivity of the 

ecosystem are likely to be compromised. Niche-based bioclimate envelope models have been widely used to project 

future geographic distributions of ecosystem climate niches. 

However, challenges arising from model accuracy as well 

as the uncertainties of future climates make it difficult to 

apply the model projections with confidence in developing 

adaptive strategies in natural resource management. The 

bioclimate envelope models are built based on the relationships between the observed presence of an ecosystem type 

and the climatic conditions of a given ecosystem. However, 

such relationships are complicated and difficult to model. 

For the future climates, there are over 140 climate change 

scenarios from the Intergovernmental Panel on Climate 

Change (IPCC) Fourth Assessment Report, and they vary 

substantially in magnitude as well as in spatial and temporal patterns. Using different climate change scenarios may 

lead to totally different adaptation strategies. Several individual scenarios can be averaged to create an “ensemble 

scenario,” but specific details are lost in this process. The 

Centre for Forest Conservation Genetics, Department of 

Forest Sciences, University of British Columbia (Tongli 

Wang and Sally Aitken), in collaboration with the Ministry 

of Forests, Lands, and Natural Resources Operations 

(Elizabeth Campbell and Greg O’Neill), have accurately 

modelled the Biogeoclimatic (BGC) ecosystem zones with 

soil sample increased during the 3 years of the study. The 

compositional similarity between seed bank and vegetation did increase for the 0–2 cm surface layer, and remained 

roughly constant for deeper soil layers. Thus, the patterns 

found differed from those of other secondary successional 

communities and do not support the tested paradigm. Rather, 

our findings resemble those from a number of primary successional ecosystems. We suggest that temporal patterns of 

similarity between seed bank and vegetation should be interpreted in terms of the factors that determine the relative rates 

of compositional change in vegetation and seed bank at different stages of succession. Our study also provides information about the seed bank persistence of 22 species, including 

several species for which such knowledge previously was 

scarce. In particular, this study indicates the extreme longevity of the seeds of the halophytic grass Puccinellia distans. 

Several years after its rapid decline in the vegetation, this 

species dominated the seed bank, and no measurable decline 

in seed density was detected during this study.

529. Wallis CM, Huber DP, Lewis KJ. 2011.

Ecosystem, location, and climate effects on foliar secondary 

metabolites of lodgepole pine populations from central British 

Columbia. Journal of Chemical Ecology 37:607-621.

Type: Journal 

Geographic Area: Western Canada

Keywords: climate change, gas chromatography-mass spectrometry (GC-MS), lignin, lodgepole pine, monoterpenoid, 

phenolics, Pinus contorta latifolia, sesquiterpenoid, diterpenoid

Abstract: Lodgepole pines, Pinus contorta Douglas ex 

Louden var. latifolia Engelm. ex S. Watson, are encountering 

increased abiotic stress and pest activity due to recent increases 

in temperature and changes in precipitation throughout their 

range. This tree species counters these threats by producing 

secondary metabolites, including phenolics and terpenoids. 

We examined foliar levels of lignin, soluble phenolics, monoterpenoids, sesquiterpenoids, and diterpenoids in 12 stands in 

British Columbia, Canada. We used these data to assess associations among foliar secondary metabolite levels and ecosystem, geographic, and climatic variables. Regressions were 

also performed to observe which combinations of variables 

best explained secondary metabolite variance. Stands of P. c. 

latifolia in the Coastal Western Hemlock and Interior Cedar/

Hemlock biogeoclimatic zones had consistently greater foliar 

levels of almost all measured secondary metabolites than did 

other stands. Lignin was present in greater amounts in Boreal 

White/Black Spruce ecosystem (i.e., northern) stands than in 

southern stands, suggesting a role for this metabolite in pine 

survival in the boreal forest. Attempts to develop regression 

models with geographic and climatic variables to explain foliar 

secondary metabolite levels resulted in multiple models with 

similar predictive capability. Since foliar secondary metabolite 

levels appeared to vary most between stand ecosystem types 

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 189

532. Wang J, Li D, Ren H, Yang L. 2010. Seed supply and the regeneration potential for plantations and shrubland in southern China. Forest Ecology and Management 

259:2390-2398.

Type: Journal 

Geographic Area: Heshan City, Guangdong, Southern China

Keywords: indigenous species, plant recruitment, seed bank, 

seed dispersal, seed limitation, succession

Abstract: Assessing the characteristics of seed supply will 

be vital to better understand the dynamics of forest regeneration. In this study, we surveyed the aboveground vegetation, 

the seed rain, the seed bank, and natural seedling emergence 

in four typical 24-year-old plantations (eucalyptus, mixednative, mixed legume, and mixed-conifer) and a naturally 

successional shrubland in southern China. The dominant 

species in the understory were similar among the five plant 

communities. The seed rain and the seed bank were dominated by shrubs and herbs but indigenous tree species were 

rare. Species that were common to all five-plant communities represented a great proportion of the seeds in the seed 

rain and seed bank. The seed rain consisted mostly of seeds 

derived from the local plant community. Seed abundance 

was greater in the seed bank than in the seed rain, and species richness was greater in the seed bank and in the corresponding plant community than in the seed rain. Species 

composition similarity between the seed rain, the seed bank, 

and the aboveground vegetation was low, because the seed 

rain contained much fewer species, and the seed bank and 

aboveground vegetation contained many different species, 

respectively. These findings indicate that both the seed rain 

and the seed bank play important roles in providing seeds 

for plant recruitment in the understory, but the seed bank 

contributes more than the current seed rain to the diversity 

of recruited plants. The current plant community has little 

impact on the qualitative composition of the seed rain and 

seed bank. Based on these data, it appears that succession 

to the desired zonal, mature forest community is unlikely 

to result from seeds in the seed rain or seed bank. Lack of 

seed availability of desired zonal mature forest species is the 

main bottleneck currently limiting succession in the plantations. Reintroduction of late-successional species could facilitate the desired succession.

533. Wang T, Campbell EM, O’Neill GA, Aitken 

SN. 2012. Projecting future distributions of ecosystem climate 

niches: uncertainties and management applications. Forest 

Ecology and Management 279:128-140.

Type: Journal 

Geographic Area: Western Canada

Keywords: climate change, forest management, ecosystem, 

climate envelope, Random Forest, consensus map

climate variables. The model was built using Random Forest 

(a machine-learning classifier) with high-resolution climate 

variables generated by ClimateWNA (http://www.genetics

.forestry.ubc.ca/cfcg/climate-models.html) and validated 

with an independent dataset. Consensus projections based 

on multiple climate change scenarios were used to cope with 

the uncertainty in future climate. The major results of this 

study are summarized below.

Link: http://www.jem.forrex.org/index.php/jem/article/viewFile/551/476

531. Wang T, O’Neill GA, Aitken SN. 2010.

Integrating environmental and genetic effects to predict responses to tree populations to climate. Ecological Applications 

20:153-163.

Type: Journal 

Geographic Area: Western Canada

Keywords: adaptation, assisted migration, British Columbia, 

Canada, ClimateBC, climate change, provenance test, response 

function, sample size, species range, transfer function

Abstract: Climate is a major environmental factor affecting 

the phenotype of trees and is also a critical agent of natural 

selection that has molded among-population genetic variation. 

Population response functions describe the environmental effect of planting site climates on the performance of a single 

population, whereas transfer functions describe among-population genetic variation molded by natural selection for climate. Although these approaches are widely used to predict 

the responses of trees to climate change, both have limitations. 

We present a novel approach that integrates both genetic and 

environmental effects into a single ‘‘universal response function’’ (URF) to better predict the influence of climate on phenotypes. Using a large lodgepole pine (Pinus contorta Dougl. 

ex Loud.) field transplant experiment composed of 140 populations planted on 62 sites to demonstrate the methodology, we 

show that the URF makes full use of data from provenance 

trials to: (1) improve predictions of climate change impacts on 

phenotypes; (2) reduce the size and cost of future provenance 

trials without compromising predictive power; (3) more fully 

exploit existing, less comprehensive provenance tests; (4) 

quantify and compare environmental and genetic effects of 

climate on population performance; and (5) predict the performance of any population growing in any climate. Finally, we 

discuss how the last attribute allows the URF to be used as a 

mechanistic model to predict population and species ranges for 

the future and to guide assisted migration of seed for reforestation, restoration, or afforestation and genetic conservation in a 

changing climate.

Link: http://izt.ciens.ucv.ve/ecologia/Archivos/ECO_

POB%202010/ECOPO2_2010/Wang%20et%20al%202010.

pdf

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190 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Abstract: Although growth response functions have previously been developed for lodgepole pine (Pinus contorta 

Dougl. ex Loud.) populations in British Columbia, new analyses were conducted: (1) to demonstrate the merit of a new local 

climate model in genecological analysis; (2) to highlight new 

methods for deriving response functions; and (3) to evaluate 

the impacts of management options for existing geographically defined seed planning units (SPUs) for reforestation. 

Results of this study suggest that new methods for anchoring 

population response functions, and a multivariate approach for 

incorporating climate variables into a single model, considerably improve the reliability of these functions. These functions 

identified a small number of populations in central areas of the 

species distribution with greater growth potential over a wide 

range of mean annual temperature (MAT). Average productivity of lodgepole pine is predicted to increase (up to 7%) if moderate warming (~2 °C MAT) occurs in the next few decades as 

predicted, although productivity would substantially decline in 

some SPUs in southern BC. Severe global warming (> 3 °C 

MAT) would result in either a drastic decline in productivity 

or local populations being extirpated in southern SPUs. New 

deployment strategies using the best seed sources for future 

reforestation may not only be able to mitigate the negative 

impact of global warming, but may even be able to increase 

productivity in some areas.

535. Webber BL, Scott JK. 2012. Rapid global change: 

implications for defining natives and aliens. Global Ecology and 

Biogeography 21:305-311.

Type: Journal 

Geographic Area: Global

Keywords: anthropogenic disturbance, assisted migration, 

climate change, dispersal, exotic, introduced, invasion, landscape modification, movement ecology

Abstract: The ability to ascribe native or alien status to species in a rapidly changing world underpins diverse research 

fields that overlap with global change and biological invasions via biodiversity. Current definitions generally link alien 

status to anthropogenic dispersal events, but this can create 

conflicts for active management and global change adaptation 

strategies, such as managed relocation and restoration ecology. Here we propose a unifying approach that allows for the 

incorporation of rapid global change into biological invasion 

terminology. We introduce the concept of a projected dispersal envelope (PDE) to define the region where a species is or 

could be native, irrespective of human involvement. The PDE 

integrates biogeography and niche theory with existing invasion terminology to place a spatial and temporal context on 

species movements. We draw on diverse suite of topical organism movements to illustrate these concepts. Our restructured 

definitions allow for native species to move into or with rapidly shifting climatic regions, as well as identifying the inappropriate introduction of alien species to new areas. Moreover, 

Abstract: Projecting future distributions of ecosystems or 

species climate niches has widely been used to assess the 

potential impacts of climate change. However, variability 

in such projections for the future periods, particularly the 

variability arising from uncertain future climates, remains 

a critical challenge for incorporating these projections into 

climate change adaptation strategies. We combined the use 

of a robust statistical modeling technique with a simple 

consensus approach consolidating projected outcomes for 

multiple climate change scenarios, and exemplify how the 

results could guide reforestation planning. Random Forest 

(RF) was used to model relationships between climate 

(1961–1990), described by 44 variables, and the geographic 

distribution of 16 major ecosystem types in British Columbia 

(BC), Canada. The model predicted current ecosystem distributions with high accuracy (mismatch rate = 4–16% for 

most ecosystem classes). It was then used to predict the 

distribution of ecosystem climate niches for the last decade 

(2001–2009) and project future distributions for 20 climate 

change scenarios. We found that geographic distributions of 

the suitable climate habitats for BC ecosystems have already 

shifted in 23% of BC since the 1970s. Consensus projections 

for future periods (2020s, 2050s, 2080s) indicated climates 

suitable for grasslands, dry forests, and moist continental 

cedar–hemlock forests would substantially expand; climate 

habitat for coastal rainforests would remain relatively stable; and habitat for boreal, subalpine and alpine ecosystems 

would decrease substantially. Using these consensus projections and data on the occurrence of Douglas-fir (Pseudotsuga 

menziesii [Mirb.] Franco) in BC ecosystems, we estimated a 

twofold increase in seedling demand for this frost-sensitive, 

commercially important timber species, suggesting managers could begin planning to expand seed inventories and 

seed orchard capacity to more widely plant this species on 

logged sites. The results of this work demonstrate the power 

of RF for building climate envelope models and illustrate the 

utility of consensus projections for incorporating uncertainty 

about future climate into management planning. It also emphasizes the immediate need for adapting natural resource 

management to a changing climate.

Link: http://bcwildfire.ca/ftp/HET/external/!publish/Web/

climate/Projecting-future-distributions-of-ecosystem-climateniches.pdf

534. Wang T, Hamann A, Yanchuk A, O’Neill GA, 

Aitken SN. 2006. Use of response functions in selecting 

lodgepole pine populations for future climates. Global Change 

Biology 12:2404-2416.

Type: Journal 

Geographic Area: Western Canada

Keywords: climate change impacts, facilitated migration, forest productivity, growth response function, local adaptation, 

Pinus contorta, seed deployment

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 191

means to preserve a species. With over half the known, wild 

federally endangered Crenulate leadplant (Amorpha herbacea 

var. crenulata), Fabaceae, growing on unprotected land slated 

for development, preserving this unprotected population was 

critical. We rescued whole plants, cuttings, and seeds for an 

experimental translocation. Into a restored pine rockland, 

once dominated by the invasive exotic tree Brazilian pepper 

(Schinus terebinthifolius), we transplanted plants from different 

sources and of different sizes. Plants used were rescued from 

an unprotected site, seedlings, and 1-, 2-, and 7-year-old plants 

from Fairchild Tropical Botanic Garden’s ex situ collection, 

creating a novel population in a new habitat. We also evaluated 

which propagule type and source had the best survival, growth, 

and reproduction. After 40 months, overall transplant survival 

was 71%. Large whole plants, rescued and nursery grown, had 

the best survival rates (86 and 78%), whereas cuttings had 67% 

survival and seedlings had only 26% survival. The restored 

site, once nearly a monoculture of S. terebinthifolius, is now 

dominated by 104 native plant species, including 17 naturally 

recruited state listed, plus the one translocated federally endangered plant species. In addition, one federally threatened snake 

species was observed on the site. These studies demonstrate 

that botanic garden collections not only play a vital role in the 

conservation of species’ genetic diversity but also can be used 

as source material for habitat restoration.

538. Williams JW, Shuman BN, Webb TI, Bartlein 

PJ, Leduc PL. 2004. Late-quaternary vegetation dynamics in North America: scaling from taxa to biomes. Ecological 

Monographs 74:309-334.

Type: Journal 

Geographic Area: North America

Keywords: biome maps, biome reconstruction, pollenbased, fossil pollen data, North America, vegetation history, 

paleoecology, plant functional types, Quaternary, vegetation 

dynamics

Abstract: This paper integrates recent efforts to map the distribution of biomes for the late Quaternary with the detailed 

evidence that plant species have responded individualistically to climate change at millennial timescales. Using a fossilpollen data set of over 700 sites, we review late-Quaternary 

vegetation history in northern and eastern North America 

across levels of ecological organization from individual taxa 

to biomes, and apply the insights gained from this review to 

critically examine the biome maps generated from the pollen data. Higher-order features of the vegetation (e.g., plant 

associations, physiognomy) emerge from individualistic 

responses of plant taxa to climate change, and different representations of vegetation history reveal different aspects of 

vegetation dynamics. Vegetation distribution and composition were relatively stable during full-glacial times (21,000–

17,000 yr BP) [calendar years] and during the mid- to late 

Holocene (7,000 – 500 yr BP), but changed rapidly during 

our definitions framework forms a timely and essential component of adaptation policies and responses for invasive species 

management and the enhancement of biodiversity in a rapidly 

changing world. 

536. Wendelberger KS, Maschinski J. 2009.

Linking geographical information systems and observational 

and experimental studies to determine optimal seedling microsites of an endangered plant in a subtropical urban fire-adapted 

ecosystem. Restoration Ecology 17:845-853.

Type: Journal 

Geographic Area: Florida, USA

Keywords: Amorpha herbacea var. crenulata, fire ecology, litter depth, management, rare plant conservation, restoration

Abstract: Understanding microsite requirements is critical 

for restoring sustainable rare plant populations and creating 

meaningful management plans that will enhance native species’ population viability. Natural areas bordered by urban 

communities have restricted size and constrained management 

options that confound goals of preserving biodiversity. Using 

geographical information systems and observational and experimental studies, we determined that current microsites 

required for seedling establishment within existing habitat 

of the federally endangered Crenulate leadplant are few and 

spatially restricted, resulting in little wild seedling recruitment. 

Experimental and wild seedlings had highest germination rates 

in litter layers 1–2 cm in depth and survived the longest in litter 0.5–2 cm deep. Fire suppression and aggressive native and 

exotic plants have accumulated litter up to 33 cm near some of 

the wild adult Crenulate leadplant. Only 9% of habitat occupied 

by this taxon at its largest population is conducive to seedling 

establishment. Similarly, 30% is conducive to seedling establishment in the second largest population. Fire plays a critical 

role in south Florida ecosystem dynamics and the restoration 

and preservation of its landscape and biodiversity. When fire 

is not possible, other management is needed. Manual litter and 

downed debris removal are recommended at both population 

sites to improve the probability of Crenulate leadplant seedling 

establishment and population persistence.

537. Wendelberger KS, Fellows MQN, Maschinski 

J. 2008. Rescue and restoration: experimental translocation 

of Amorpha herbacea Walter var. crenulata (Rybd.) Isley into a 

novel urban habitat. Restoration Ecology 16:542-552.

Type: Journal 

Geographic Area: Florida, USA

Keywords: Amorpha herbacea var. crenulata, propagule type, 

rare plant conservation, restoration, translocation

Abstract: Though translocations of rare populations should 

be considered only as the last resort for species’ conservation, when habitat destruction is imminent, it may be the only 

CHAPTER 3 • MIGRATION • RESEARCH
192 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

half of forestland densities with the majority of urban tree 

species found in nearby (< 100 km) forestland. Urban tree 

propagation is often facilitated by humans, whereas the necessary pollinators and agents of tree seed dispersal in forestlands may be lacking regardless of climate change. It is 

suggested that urban areas may serve divergent, dual roles 

as both a native tree seed source and refuge for a limited 

number of forestland tree species, but also a facilitator of 

non-native tree invasion.

Link: http://www.treesearch.fs.fed.us/pubs/34734

540. Zhu K, Woodall CW, Clark JS. 2012. Failure 

to migrate: lack of tree range expansion in response to climate 

change. Global Change Biology 18:1042-1052.

Type: Journal 

Geographic Area: Eastern USA

Keywords: biogeography, climate change, Forest Inventory 

and Analysis, latitude, presence/absence, range shift, seedling, 

tree migration

Abstract: Tree species are expected to track warming climate by shifting their ranges to higher latitudes or elevations, but current evidence of latitudinal ranges shifts for 

suites of species is largely indirect. In response to global 

warming, off-spring of trees are predicted to have ranges 

extend beyond adults at leading edges and the opposite relationship at trailing edges. Large-scale forest inventory data 

provide an opportunity to compare present latitudes of seedlings and adult trees at their range limits. Using the USDA 

Forest Service’s Forest Inventory and Analysis data, we directly compared seedling and tree 5th and 95th percentile 

latitudes for 92 species in 30 longitudinal bands for 43,334 

plots across the eastern United States. We further compared 

these latitudes with 20th century temperature and precipitation change and functional traits, including seed size and 

seed spread rate. Results suggest that 58.7% of the trees species examined show the pattern expected for a population 

undergoing range contraction, rather than expansion, at both 

northern and southern boundaries. Fewer species show a 

pattern consistent with a northward shift (20.7%) and fewer 

still with a southward shift (16.3%). Only 4.3% are consistent with expansion at both range limits. When compared 

with the 20th century climate changes that have occurred 

at the range boundaries themselves, there is no consistent 

evidence that population spread is greatest in areas where 

climate has changed most; nor are patterns related to seed 

size or dispersal characteristics. The fact that the majority of 

seedling extreme latitudes are less than those for adult trees 

may emphasize the lack of evidence for climate-mediated 

migration, and should increase concerns for the risks posed 

by climate change. 

Link: http://www.nicholas.duke.edu/people/faculty/clark/

pages/publications/gcbZhu2011.pdf

the late-glacial period and early Holocene (16,000 – 8,000 yr 

BP) and after 500 yr BP. Shifts in plant taxon distributions 

were characterized by individualistic changes in population 

abundances and ranges and included large east–west shifts 

in distribution in addition to the northward redistribution of 

most taxa. Modern associations such as Fagus–Tsuga and 

Picea–Alnus–Betula date to the early Holocene, whereas 

other associations common to the late-glacial period (e.g., 

Picea–Cyperaceae–Fraxinus–Ostrya/Carpinus) no longer exist. Biomes are dynamic entities that have changed in distribution, composition, and structure over time. The late-Pleistocene suite of biomes is distinct from those that grew during 

the Holocene. The pollen-based biome reconstructions are 

able to capture the major features of late-Quaternary vegetation but downplay the magnitude and variety of vegetational 

responses to climate change by (1) limiting apparent landcover change to ecotones, (2) masking internal variations in 

biome composition, and (3) obscuring the range shifts and 

changes in abundance among individual taxa. The compositional and structural differences between full-glacial and 

recent biomes of the same type are similar to or greater than 

the spatial heterogeneity in the composition and structure of 

present-day biomes. This spatial and temporal heterogeneity 

allows biome maps to accommodate individualistic behavior 

among species but masks climatically important variations 

in taxonomic composition as well as structural differences 

between modern biomes and their ancient counterparts.

Link: http://www.clas.ufl.edu/users/mbinford/GEOXXXX_

Biogeography/LiteratureForLinks/Williams_et_al_2004_Late_

Quaternary_veg_dynamics-scaling_taxa_to_biomes_EcolMonogr.pdf

539. Woodall CW, Nowak DJ, Liknes GC, Westfall 

JA. 2010. Assessing the potential for urban trees to facilitate 

forest tree migration in the eastern United States. Forest Ecology 

and Management 259:1447-1454.

Type: Journal 

Geographic Area: Eastern USA

Keywords: climate change, urban, facilitated migration, assisted migration, eastern United States, forest ecosystems

Abstract: Latitudinal shifts in tree species distributions are 

a potential impact of climate change on forest ecosystems. 

It has been hypothesized that some tree species may become 

extirpated as climate change effects may exceed their migration ability. The goal of this study was to compare tree 

species compositions in northern urban areas to tree compositions in forestland areas in the eastern U.S. as an indicator 

of the potential for urban trees to facilitate future forest tree 

species migration. Results indicated that a number of tree 

species native to eastern U.S. forests of southern latitudes 

are currently present in northern urban forests. The biomass 

density (Mg/ha) of urban tree species is typically less than 
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 193

541. Bergelson J, Roux F. 2010. Towards identifying 

genes underlying ecologically relevant traits in Arabidopsis thaliana. Nature Reviews Genetics 11:867-879.

Type: Journal 

Geographic Area: France, Europe

Compilers’ Keywords: biological adaptation, Arabidopsis 

thaliana, ecosystem, environment, plant genes, genetic variation, genome-wide association study, review

Abstract: A major challenge in evolutionary biology and 

plant breeding is to identify the genetic basis of complex 

quantitative traits, including those that contribute to adaptive 

variation. Here we review the development of new methods 

and resources to fine-map intraspecific genetic variation that 

underlies natural phenotypic variation in plants. In particular, 

the analysis of 107 quantitative traits reported in the first genome-wide association mapping study in Arabidopsis thaliana

sets the stage for an exciting time in our understanding of plant 

adaptation. We also argue for the need to place phenotypegenotype association studies in an ecological context if one is 

to predict the evolutionary trajectories of plant species.

Link: http://www.ncbi.nlm.nih.gov/pubmed/21085205

542. Booth TH, Williams KJ, Belbin L. 2012.

Developing biodiverse plantings suitable for changing climatic 

conditions 2: using the Atlas of Living Australia. Ecological 

Management & Restoration 13:274-281.

Type: Journal 

Geographic Area: Australia

Keywords: Atlas of Living Australia, biodiversity, climate 

change, revegetation, maxent models, species distribution 

models

Abstract: There has been an increasing investment of taxpayer dollars in revegetation in Australia over the past 20 years, at 

both federal and state levels. The largest of these, the Australian 

Government’s Biodiversity Fund, will invest A$946 million to 

revegetate, rehabilitate and restore landscapes to store carbon, 

enhance biodiversity and build environmental resilience under 

climate change. The universal challenge for restoration practitioners working within these programmes is species selection 

for both current and future environmental conditions at a given 

site. For policy makers, the challenge is to provide guidelines 

and tools for this process. The first paper in this series of two 

papers looked at scientific methods that could provide underpinning knowledge to improve the assessment of species 

vulnerability to climatic and atmospheric change. In this paper, the publically accessible Atlas of Living Australia is used 

to demonstrate how revegetation project leaders can assess 

General (339–380)—debates, reviews, 

trends, surveys, summaries

Research (381–540)—migration studies, 

range shifts, reintroductions

Strategies (541–580)—adaptation options, 

frameworks, decision support

Resources (581–582)—tools, websites, software

Migration

CHAPTER 3 • MIGRATION • STRATEGIES

#. Author(s), Date. Title. Source.

Type: 

Geographic Area: 

Keywords: 

Description:

Link:

location of study

keywords written by the 

author(s) or provided by the 

compilers

ID number and

information

abstract, summary, or 

portions of the introduction 

or conclusion written by the 

author(s) or provided by the 

compilers

book, book section, conference 

paper, dissertation, government 

document, journal article, 

newsletter, presentation, report, 

thesis or web page

internet address of reference

Each reference contains the following:
194 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

local adaptations of foundation or keystone species, and that 

these local ecotypes should be moved within their own range. 

Hence, this type of relocation could be applied with minimal 

risk in many restoration efforts. We think that caution is needed 

when considering the translocation of these foundation species, even within their range. Many recent studies have shown 

that foundation species can influence community structure and 

ecosystem processes through heritable traits, which suggests 

a genetic basis for ecosystem services. Thus, the translocation 

of different genotypes of foundation species might lead to unexpected results of colonization and might not be as “predictable” as Kreyling et al. have argued. Here, in our response, we 

stress how AC of foundation species can have important evolutionary consequences that might be impossible to reverse. 

We propose, whenever possible, 1) to favor population mixes 

of the foundation species to minimize the potential negative 

effects of specific genotypes; and 2) to collect from adjacent 

populations along ecological clines of the foundation species 

to mimic natural processes of migration under climate change. 

Link: http://www.researchgate.net/publication/233207938_

Assisted_Colonization_of_Foundation_Species_Lack_of_

Consideration_of_the_Extended_Phenotype_ConceptResponse_to_Kreyling_et_al._(2011)/file/9fcfd509c2919ebb6f.

pdf

545. Garcia D, Zamora R, Amico GC. 2010. Birds 

as suppliers of seed dispersal in temperate ecosystems: conservation guidelines from real-world landscapes. Conservation 

Biology 24:1070-1079.

Type: Journal 

Geographic Area: Global

Keywords: anthropogenic landscapes, Cantabrian forest, 

ecosystem services, fleshy-fruited plants, frugivorous birds, 

Mediterranean shrubland, mobile links, Patagonian forest, 

seed dispersal

Abstract: Seed dispersal by animals is considered a pivotal 

ecosystem function that drives plant-community dynamics 

in natural habitats and vegetation recovery in human-altered 

landscapes. Nevertheless, there is a lack of suitable ecological knowledge to develop basic conservation and management 

guidelines for this ecosystem service. Essential questions, such 

as how well the abundance of frugivorous animals predicts 

seeding function in different ecosystems and how anthropogenic landscape heterogeneity conditions the role of dispersers, remain poorly answered. In three temperate ecosystems, 

we studied seed dispersal by frugivorous birds in landscape 

mosaics shaped by human disturbance. By applying a standardized design across systems, we related the frequency of 

occurrence of bird-dispersed seeds throughout the landscape 

to the abundance of birds, the habitat features, and the abundance of fleshy fruits. Abundance of frugivorous birds in itself predicted the occurrence of dispersed seeds throughout 

whether the species and provenances used in their revegetation 

projects are likely to be suitable for changing environmental 

conditions. While using the Atlas can assist current selections, 

ways in which more reliable selections for changing climatic 

conditions could be made are also outlined.

543. Chauvenet ALM, Ewen JG, Armstrong DP, 

Blackburn TM, Pettorelli N, Gompper M. 2012.

Maximizing the success of assisted colonizations. Animal 

Conservation 16:161-169.

Type: Journal 

Geographic Area: Global

Keywords: translocation, climate-change adaptation, population dynamics, species distribution modelling, assisted migration, managed relocation

Abstract: Climate change is causing spatio-temporal shifts in 

environmental conditions, and species that are not able to track 

suitable environments may face increased risks of extinction. 

Assisted colonization, a form of translocation, has been proposed as a tool to help species survive the impacts of climate 

change. Unfortunately, translocations generally have a low 

success rate, a well-documented fact that is not considered in 

most of the recent literature on assisted colonization. One of 

the main impediments of translocation success is inadequate 

planning. In this review, we argue that by using well-known 

analytical tools such as species distribution models and population dynamics modelling we can maximize the success of 

assisted colonization. In particular, we present guidelines as to 

which questions should be investigated when planning assisted 

colonization and suggest methods for answering them. Finally, 

we also highlight further implementation and research issues 

that remain to be solved for assisted colonizations to be efficient climate change adaptation tools. 

Link: http://www.researchgate.net/publication/233895747_

Maximizing_the_success_of_assisted_colonizations/

file/79e4150d036521d74c.pdf

544. Frascaria-Lacoste N, Fernández-Manjarrés 

J. 2012. Assisted colonization of foundation species: lack of 

consideration of the extended phenotype concept-response to 

Kreyling et al. (2011). Restoration Ecology 20:296-298.

Type: Journal 

Geographic Area: Global

Keywords: assisted migration, ecosystem engineers, forest tree, genetic diversity, managed translocation, restoration 

actions

Abstract: Assisted colonization (AC), or the intentional 

translocation of populations to compensate for risks related to 

climate change, is receiving increasing attention. It has been 

recently suggested by Kreyling et al. (2011) that rather than 

relocating endangered species, a focus should be placed on 

CHAPTER 3 • MIGRATION • STRATEGIES
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 195

conform well to projections for the 2020s, which suggests that 

it is a safe strategy to change current reforestation practices 

and adapt to new climatic realities through assisted migration 

prescriptions.

Link: http://ualberta.ca/~ahamann/publications/pdfs/Gray_

Hamann_2011.pdf

547. Hannah L, Midgley GF, Andelman S, Araujo 

M, Hughes G, Martinez-Meyer E, Pearson R, 

Williams P. 2007. Protected area needs in a changing climate. Frontiers in Ecology and the Environment 5:131-138.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: range shifts, conservation, species 

distribution modeling, biodiversity

Abstract: Range shifts due to climate change may cause 

species to move out of protected areas. Climate change could 

therefore result in species range dynamics that reduce the relevance of current fixed protected areas in future conservation 

strategies. Here, we apply species distribution modeling and 

conservation planning tools in three regions (Mexico, the Cape 

Floristic Region of South Africa, and Western Europe) to examine the need for additional protected areas in light of anticipated species range shifts caused by climate change. We set 

species representation targets and assessed the area required 

to meet those targets in the present and in the future, under a 

moderate climate change scenario. Our findings indicate that 

protected areas can be an important conservation strategy in 

such a scenario, and that early action may be both more effective and less costly than inaction or delayed action. According 

to our projections, costs may vary among regions and none of 

the three areas studied will fully meet all conservation targets, 

even under a moderate climate change scenario. This suggests 

that limiting climate change is an essential complement to adding protected areas for conservation of biodiversity.

Link: http://courses.washington.edu/cfr550/pdfs/Hannah%20

et%20al%202007.pdf

548. Havens K, Vitt P, Still S, Kramer AT, Fant JB, 

Schatz K. 2015. Seed sourcing for restoration in an era of 

climate change. Natural Areas Journal 35:122-133.

Type: Journal

Geographic Area: USA

Keywords: assisted migration, climate change, provenance, 

restoration, seed sourcing

Abstract: Proper sourcing of seed for ecological restoration 

has never been straightforward, and it is becoming even more 

challenging and complex as the climate changes. For decades, 

restoration practitioners have subscribed to the “local is best” 

tenet, even if the definition of “local” was often widely divergent between projects. However, given our increasing ability 

the landscape in all ecosystems studied. Even those landscape 

patches impoverished due to anthropogenic disturbance received some dispersed seeds when visited intensively by birds. 

Nonetheless, human-caused landscape degradation largely affected seed-deposition patterns by decreasing cover of woody 

vegetation or availability of fruit resources that attracted birds 

and promoted seed dispersal. The relative role of woody cover 

and fruit availability in seed dispersal by birds differed among 

ecosystems. Our results suggest that to manage seed dispersal 

for temperate ecosystem preservation or restoration one should 

consider abundance of frugivorous birds as a surrogate of landscape-scale seed dispersal and an indicator of patch quality for 

the dispersal function; woody cover and fruit resource availability as key landscape features that drive seedfall patterns; 

and birds as mobile links that connect landscape patches of 

different degrees of degradation and habitat quality via seed 

deposition.

Link: http://www.ncbi.nlm.nih.gov/pubmed/20136873

546. Gray LK, Hamann A. 2011. Strategies for reforestation under uncertain future climates: guidelines for Alberta, 

Canada. PLoS ONE 6:1-9.

Type: Journal 

Geographic Area: Alberta, Canada

Compilers’ Keywords: climate change, commercial forestry, 

assisted migration, projections, distribution

Abstract: Commercial forestry programs normally use locally collected seed for reforestation under the assumption that 

tree populations are optimally adapted to local environments. 

However, in western Canada this assumption is no longer valid 

because of climate trends that have occurred over the last several decades. The objective of this study is to show how we can 

arrive at reforestation recommendations with alternative species and genotypes that are viable under a majority of climate 

change scenarios. In a case study for commercially important 

tree species of Alberta, we use an ecosystem based bioclimate 

envelope modeling approach for western North America to 

project habitat for locally adapted populations of tree species 

using multi-model climate projections for the 2020s, 2050s 

and 2080s. We find that genotypes of species that are adapted 

to drier climatic conditions will be the preferred planting stock 

over much of the boreal forest that is commercially managed. 

Interestingly, no alternative species that are currently not present in Alberta can be recommended with any confidence. 

Finally, we observe large uncertainties in projections of suitable habitat that make reforestation planning beyond the 2050s 

difficult for most species. More than 50,000 hectares of forests 

are commercially planted every year in Alberta. Choosing alternative planting stock, suitable for expected future climates, 

could therefore offer an effective climate change adaptation 

strategy at little additional cost. Habitat projections for locally adapted tree populations under observed climate change 

CHAPTER 3 • MIGRATION • STRATEGIES
196 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Geographic Area: USA

Compilers’ Keywords: policy, forest management, species 

rescue, ecosystem

Introduction (5th paragraph): Actions associated with 

assisted migration cover a wide variety of movements for a 

number of different purposes, ranging from moving a seed 

source to another location within the species range in order 

to maintain ecosystem productivity, to moving a suite of species, or a community, outside of its historical range to prevent 

extinction. This paper examines two major categories of assisted migration that primarily impact management decisions 

on National Forest System (NFS) lands, assisted migration 

to maintain ecosystem services (Ecosystem AM) and species 

rescue assisted migration (Species Rescue AM); these definitions closely follow what is presented by Pedlar et al. (2012) 

for forest trees. These two categories differ in the types of species managed, management objectives, relative feasibility, and 

associated risks. Most discussions in the literature focus on 

species rescue and little on Ecosystem AM. 

Link: http://www.treesearch.fs.fed.us/pubs/45424

551. Jones TA, Monaco TA. 2009.A role for assisted evolution in designing native plant materials for domesticated landscapes. Frontiers in Ecology and the Environment 7:541-547.

Type: Journal 

Geographic Area: Western USA

Compilers’ Keywords: restoration, assisted migration, 

genetic patterns, function, concepts, native plant material 

development

Abstract: Developers of native plant propagation materials 

for wildland restoration may emphasize naturally occurring 

genetic patterns or, in contrast, the material’s empirical performance in comparative field trials. We contend that both 

approaches have value and need not be mutually exclusive. 

Anthropogenic influences have pushed many ecosystems 

across ecological thresholds, to less desirable states, so that actively managing for “domesticated nature”—nature as modified, either intentionally or inadvertently, by humans—is more 

realistic and more likely to succeed than recreating the original 

ecosystem. Furthermore, when domesticated nature is the most 

reasonable objective, empirical performance, together with 

geographical origin, are plausible criteria for choosing restoration plant material. For altered ecosystems, we suggest that 

evolution should be assisted by the inclusion of plants that (1) 

reflect general historical evolutionary patterns, (2) are particularly suited to the modified environment, (3) are able to adapt 

to contemporary selection pressures, and (4) contribute to the 

restoration of ecosystem structure and function.

Link: http://sfc.smallfarmcentral.com/dynamic_content/

uploadfiles/152/jones-a%20role%20for%20assisted%20evolution.pdf

to characterize habitats, and rapid climate change, we can no 

longer assume that locally sourced seeds are always the best or 

even an appropriate option. We discuss how plants are responding to changing climates through plasticity, adaptation, and migration, and how this may influence seed sourcing decisions. 

We recommend focusing on developing adequate supplies of 

“workhorse” species, undertaking more focused collections in 

both “bad” years and “bad” sites to maximize the potential to 

be able to adapt to extreme conditions as well as overall genetic diversity, and increasing seed storage capacity to ensure 

we have seed available as we continue to conduct research to 

determine how best to deploy it in a changing climate.

Link: http://www.bioone.org/doi/abs/10.3375/043.035.0116

549. Hoegh-Guldberg O, Hughes L, McIntyre S, 

Lindenmayer DB, Parmesan C, Possingham HP, 

Thomas CD. 2008. Assisted colonization and rapid climate 

change. Science 321:345-346.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: conservation, extinction risk, decision framework, translocation, assisted migration, policy forum, review, tool 

Synopsis (4th paragraph): We developed a decision framework that can be used to outline potential actions under a suite 

of possible future climate scenarios. Determining whether a 

species faces significant risk of decline or extinction under climate change requires an in-depth knowledge of the underlying 

species’ biology as well as the biological, physical, and chemical changes occurring within its environment. The risk of extinction for many widespread, generalist species found across a 

range of habitats may be low. In this case, the option of moving 

such species outside their present ranges would be dismissed. 

Some species will also disperse sufficiently to maintain large 

populations and range sizes (for example, highly dispersive 

insects or birds with generalist life histories) and others may 

adapt in situ. Where species are perceived as being at moderate risk from climate change, improvements in connectivity to 

actual or potential habitat at higher latitudes and altitudes may 

be sufficient.

Link: http://www.law.arizona.edu/depts/ele/AdaptationConference/PDFs/ParmesanAssistedColonization.pdf

550. Johnson R, Boyce S, Brandt L, Erickson VJ, 

Iverson LR, Kujawa G, Stritch L, Tkacz B. 2013.

Policy and strategy considerations for assisted migration on 

USDA Forest Service Lands. In: Browing J, Palacios P, editors. 

Proceedings of the 60th Annual Western International Forest 

Disease Work Conference. Tahoe City (CA): USDA Forest 

Service. p 35-41.

Type: Conference Paper

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 197

Keywords: forestry, climate change, policy development, 

discourse analysis, new institutionalism, forest genetics 

conservation

Abstract: The idea of moving species beyond their historic 

range, known as assisted migration, challenges some of the 

conservation ideas and norms guiding sustainable forest management in North America: relying on historical benchmarks, 

mimicking natural disturbances, and reproducing current ecological species assemblages. In this context, our paper focuses 

on how an assisted migration policy for western larch has been 

developed in British Columbia, Canada, specifically examining institutional change at three different levels of policy ideas 

and in discourses oriented internally or externally in the policy 

process. Based on 46 interviews with policy actors across 

Canada, our results suggest that the deployment of the first 

assisted migration policy in Canada successfully avoided the 

controversy surrounding the idea in the scientific community 

by changing the scientific discourse associated with best forest 

management practices. The shift from an ecological discourse 

to a genetics discourse over forest policy in British Columbia 

signals what we might expect in future forest adaptation policy 

development in Canada.

Link: http://www.sciencedirect.com/science/article/pii/

S0959378014002003

554. Koralewski TE, Wang H-H, Grant WE, Byram 

TD. 2015. Plants on the move: assisted migration of forest trees 

in the face of climate change. Forest Ecology and Management 

344:30-37.

Type: Journal 

Geographic Area: Southeastern USA

Keywords: assisted migration, Categorical Universal 

Response Function, climate change, Pinus taeda, seed deployment, Universal Response Function

Abstract: Climate change is impacting distributions of both 

fauna and flora, including many crops. In order to ensure the 

health and vigor of the species we depend upon for food and fiber, assisted migration strategies may need to be implemented. 

This is especially true for long-lived forest trees. Multinomial 

logit regression was used to develop a Categorical Universal 

Response Function (CURF) to delineate deployment zones 

for loblolly pine (Pinus taeda L.) using 15-year measurements 

from the Western Gulf Forest Tree Improvement Program 

(WGFTIP) Geographic Seed Source Study (GSSS). The 

CURF uses performance categories for the response variable, 

and the model assigns the probabilities with which the performance for a given seed source will fall into these various 

categories. First and second powers of minimum temperature 

of the coldest month, summer precipitation, and variation 

measures of these two metrics at both the seed source site of 

origin and the test location were used as independent variables. 

Planted tree volume, accounting for both survival and growth, 

552. Klenk NL. 2015. The development of assisted migration policy in Canada: an analysis of the politics of composing 

future forests. Land Use Policy 44:101-109.

Type: Journal 

Geographic Area: Canada

Keywords: discursive institutionalism, forestry, climate 

change, policy development, actor-network theory, agonistic 

pluralism

Abstract: One could argue that the development of western 

larch assisted migration (AM) policy in British Columbia 

was the result of a “policy window” largely caused by factors external to the actions of policy actors: western larch 

plantations in northern BC indicate that the species thrives 

even if planted far beyond its current range, thus representing an economic motive for AM; climate change projections 

and the mountain pine beetle epidemic represented a crisis 

situation that facilitated the deployment of a new and controversial policy option, the assisted migration of western larch, 

in forest policy in British Columbia. However, this “policy 

window” explanation disregard the relationship between the 

performative meaning of AM in its social and ecological 

context, and masks the actual politics of the discursive practices enlisting particular actor-networks in the western larch 

AM policy process. My analysis suggests that the western 

larch AM policy emerged from the relationship of specific 

policy actors with specific non-humans actors—including 

exotic and invasive species. Yet, aware of potential political 

risks of deploying this controversial practice, policy actors 

in British Columbia carefully circumscribed the purpose of 

western larch AM, distancing the western larch AM policy 

from exotic and invasive species. Additionally, western 

larch AM was framed as a natural extension of current tree 

regeneration standards and best practices in BC—thus seemingly representing “business as usual”, rather than a major 

reassemblage of the actor-networks structuring forest policy 

in BC. Comparing western larch policy discourses to policy 

discourses on AM in other provinces suggest that the differences in what actor networks are enlisted largely explains 

why no other province in Canada has yet to develop an assisted migration policy. The policy implications of this study 

are that in the composition of Canada’s future forests we 

should keep exotic and invasive species present in view.

Link: http://www.sciencedirect.com/science/article/pii/

S0264837714002749

553. Klenk NL, Larson BMH. 2015. The assisted migration of western larch in British Columbia: a signal of institutional 

change in forestry in Canada? Global Environmental Change 

31:20-27.

Type: Journal 

Geographic Area: Western Canada, British Columbia

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198 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

perceived by some as a last option for conserving endangered 

species and by others as a risky and unwise management effort 

due to current gaps of knowledge. Based on the pros and cons 

of the recent debate, we show that the current discussion mainly focuses on the assisted colonization of rare and endangered 

species beyond their natural range of distribution. We suggest 

that a more useful approach for the conservation of endangered 

species could occur by focusing on the relevant foundation or 

keystone species, which ensure ecosystem integrity for a multitude of dependent species by governing the habitat structure 

and microclimate of the site. Examples of foundation species 

include dominant tree species in forests or dominant corals in 

coral reefs. For a given conservation or restoration need (e.g. 

conservation of rare species), we recommend the assisted 

colonization of pre-adapted ecotypes of the relevant foundation species from climates similar to future expectations for the 

target site. This approach could lead to climate safe habitats for 

endangered species with minimal adverse effects on recipient 

ecosystems.

Link: http://www.researchgate.net/publication/229953735_

Assisted_Colonization_A_Question_of_Focal_Units_and_

Recipient_Localities/file/e0b4951a5c24071def.pdf

557. Krosby M, Tewksbury J, Haddad NM, 

Hoekstra J. 2010. Ecological connectivity for a changing 

climate. Conservation Biology 24:1686-1689.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: biodiversity, climate change, conservation, ecosystem, migration, 

Introduction (1st paragraph): A frequently proposed 

strategy to reduce the negative effects of climate change on 

biological diversity is to increase ecological connectivity the 

flow of organisms and ecological processes across landscapes. 

Traditionally, conservation professionals have sought to maintain or restore connectivity to ensure gene flow among isolated 

populations and promote recolonization of vacant patches. 

Given the rapid emergence of connectivity enhancement as 

a climate-change adaptation strategy, we considered whether 

connectivity should be emphasized in conservation strategies 

as global or regional temperatures increase and what principles 

for connectivity enhancement could be applied to maximize 

the usefulness of the strategy.

Link: http://www.ncbi.nlm.nih.gov/pubmed/20961330

558. Leites LP, Rehfeldt GE, Robinson AP, 

Crookston NL, Jaquish B. 2012. Possibilities and 

limitations of using historic provenance tests to infer forest 

species growth responses to climate change. Natural Resource 

Modeling 25:409-433.

Type: Journal 

was used as the response variable. Model performance was 

good, with the AUC score ranging from 0.785 to 0.808, depending on (1) whether or not the variable interactions were 

included and (2) the variable selection criterion used (AIC or 

BIC). Resulting models were then applied to historic weather 

patterns to illustrate inferred deployment zones for three seed 

sources. The projected performance generally agreed with the 

current consensus on loblolly pine seed movement guidelines. 

The models developed here can be readily implemented in a 

Decision Support System as they (1) suggest sets of adapted 

loblolly pine families from which foresters can choose based 

on local knowledge, (2) can be easily expanded to include other variables, and (3) can be applied to outputs from projected 

climate scenarios to extrapolate into the future.

Link: http://www.sciencedirect.com/science/article/pii/

S0378112715000729

555. Kostyack J, Lawler JJ, Goble DD, Olden JD, 

Scott JM. 2011. Beyond reserves and corridors: policy solutions to facilitate the movement of plants and animals in a changing climate. BioScience 61:713-719.

Type: Journal 

Geographic Area: USA

Keywords: climate change, adaptation, species movement, 

policy

Abstract: As the Earth’s climate changes, many species will 

have to move across human-dominated landscapes to track 

suitable climates and changing ecosystems. Given the magnitude of projected future climate change, expanding and 

connecting reserve networks—two of the most commonly 

recommended adaptation strategies for protecting biodiversity 

in a changing climate—will be necessary but insufficient for 

preventing climate-induced extinctions. In the present , we 

explore additional policy options that could be implemented 

to facilitate species movements in a changing climate. We discuss both existing and new policies that have the potential to 

increase landscape permeability, protect species on the move, 

and physically move species to address climate change.

556. Kreyling J, Bittner T, Jaeschke A, Jentsch A, 

Jonas Steinbauer M, Thiel D, Beierkuhnlein C. 

2011. Assisted colonization: a question of focal units and recipient localities. Restoration Ecology 19:433-440.

Type: Journal 

Geographic Area: Global

Keywords: adaptation strategy, assisted migration, core species, ecosystem engineers, managed relocation, managed 

translocation, species conservation

Abstract: Assisted colonization as an adaptation strategy to 

conserve or restore biodiversity in the face of climate change 

deservedly evokes controversy. Assisted colonization is 

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assisted colonization and sought to identify other conservation 

techniques that also display potential to promote the colonization and adaptation of species in response to climate change. 

We propose an integrated conservation strategy that includes 

management for habitat connectivity, conservation genetics, 

and when necessary, assisted colonization of species that are 

still unable to shift their ranges even given implementation of 

the above standard conservation approaches. We argue that 

this integrated approach will facilitate persistence for a larger 

proportion of species than is possible by solely using assisted 

colonization. Furthermore, a multi-faceted approach will likely 

reduce the uncertainty of conservation outcomes and will become increasingly necessary for conservation of biodiversity 

in a changing climate.

560. Lunt ID, Byrne M, Hellmann JJ, Mitchell NJ, 

Garnett ST, Hayward MW, Martin TG, McDonaldMaddden E, Williams SE, Zander KK. 2013. Using 

assisted colonization to conserve biodiversity and restore ecosystem function under climate change. Biological Conservation 

157:172-177.

Type: Journal 

Geographic Area: Global

Keywords: ecological replacement, managed relocation, climate change adaptation, ecosystem management, restoration, 

translocation

Abstract: Assisted colonization has received considerable 

attention recently, and the risks and benefits of introducing 

taxa to sites beyond their historical range have been vigorously debated. The debate has primarily focused on using 

assisted colonization to enhance the persistence of taxa 

that would otherwise be stranded in unsuitable habitat as a 

consequence of anthropogenic climate change and habitat 

fragmentation. However, a complementary motivation for 

assisted colonization could be to relocate taxa to restore 

declining ecosystem processes that support biodiversity in 

recipient sites. We compare the benefits and risks of species 

introductions motivated by either goal, which we respectively term ‘push’ versus ‘pull’ strategies for introductions 

to preserve single species or for restoration of ecological 

processes. We highlight that, by focusing on push and neglecting pull options, ecologists have greatly under estimated potential benefits and risks that may result from assisted 

colonization. Assisted colonization may receive higher 

priority in climate change adaptation strategies if relocated 

taxa perform valuable ecological functions (pull) rather than 

have little collateral benefit (push). Potential roles include 

enhancing resistance to invasion by undesired species, supporting co-dependent species, performing keystone functions, providing temporally critical resources, replacing taxa 

of low ecological redundancy, and avoiding time lags in the 

provisioning of desired functions.

Geographic Area: North America

Keywords: climate-change response functions, provenance 

tests, genotype by environment interaction, provenance transfer functions, Larix occidentalis Nutt, linear mixed-effects 

models

Abstract: Under projected changes in global climate, the 

growth and survival of existing forests will depend on their 

ability to adjust physiologically in response to environmental change. Quantifying their capacity to adjust and whether 

the response is species- or population-specific is important 

to guide forest management strategies. New analyses of 

historic provenance tests data are yielding relevant insights 

about these responses. Yet, differences between the objectives used to design the experiments and current objectives 

impose limitations to what can be learned from them. Our 

objectives are (i) to discuss the possibilities and limitations 

of using such data to quantify growth responses to changes in 

climate and (ii) to present a modeling approach that creates 

a species- and population-specific model. We illustrate the 

modeling approach for Larix occidentalis Nutt. We conclude 

that the reanalysis of historic provenance tests data can lead 

to the identification of species that have population-specific 

growth responses to changes in climate, provide estimates of 

optimum transfer distance for populations and species, and 

provide estimates of growth changes under different climate 

change scenarios. Using mixed-effects modeling techniques 

is a sound statistical approach to overcome some of the limitations of the data.

Link: http://www.treesearch.fs.fed.us/pubs/41324

559. Loss SR, Terwilliger LA, Peterson AC. 2011.

Assisted colonization: integrating conservation strategies in the 

face of climate change. Biological Conservation 144:92-100.

Type: Journal 

Geographic Area: Global

Keywords: assisted colonization, assisted migration, managed relocation, climate change, landscape connectivity, conservation genetics

Abstract: Global climate change poses an immense challenge for conservation biologists seeking to mitigate impacts 

to species and ecosystems. Species persistence will depend on 

geographic range shifts or adaptation in response to warming 

patterns as novel climates and community assemblages arise. 

Assisted colonization has been proposed as a method for addressing these challenges. This technique, which consists of 

transporting species to a new range that is predicted to be 

favorable for persistence under future climate scenarios, has 

become the subject of controversy and discussion in the conservation community due to its highly manipulative nature, 

questions about widespread feasibility, and uncertainty associated with the likelihood of translocated species becoming invasive. We reviewed the discussion and criticism associated with 

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200 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

characteristic of the species. At the southern limits of distribution, growth and competitive ability of the species will decline, 

leading to successional changes.

Link: http://treephys.oxfordjournals.org/content/14/7-8-9/797.full.pdf

563. Matyas C. 1996. Climatic adaptation of trees: 

rediscovering provenance tests. Euphytica 92:45-54.

Type: Journal 

Geographic Area: Global

Keywords: adaptation, climate change, provenance research, 

phenotypic stability, natural selection, genetic diversity

Summary: Common garden testing of populations of different origin started with forest trees more than two hundred years 

ago. Since then, so-called provenance tests have been established with most commercially important species. Beyond the 

strictly silvicultural goals, the tests offer excellent opportunities to study intraspecific genetic variation patterns and represent probably the most powerful available tool for testing 

hypotheses of climatic adaptation in trees. Analysis of adaptive 

traits (mostly juvenile height growth) in provenance experiments indicate the existence of very effective constraints on 

adaptedness. The performance of populations plotted against 

an ecological-climatic factor exhibits a characteristic pattern 

and can be described by response functions. The population 

average of a fitness related trait for a locally adapted population is often significantly lower than that of populations from 

other environments; usually the ones from milder climate 

perform better. The phenomenon is interpreted as adaptation 

lag. Suboptimal adaptation is compensated by a high level of 

genetic diversity. Molecular genetic studies confirm the high 

level of allelic and individual genetic diversity in forest trees. 

A consequence of individual homeostasis, phenotypic stability 

of populations is usually also high; the sensitivity to environmental changes is generally moderate. Phenotypically stable 

populations are valuable not only because of a wider range of 

potential cultivation but specifically because of a greater ability 

to adjust to unexpected changes. This trait should receive more 

attention in the future for obvious reasons. The maintenance 

of a high within-population genetic variance is favored by 

the genetic system of the investigated species (effective gene 

flow, outbreeding, high genetic load, etc.). Random events and 

long-lasting biotic interactions are further effects impairing 

the efficiency of natural selection. In view of expected climate 

instability, genetic adaptability of forest trees causes serious 

concern due to their long lifespan compared to the rapidity of 

expected changes in environmental conditions. The potential 

of provenance tests to interpret long-term adaptation processes 

should be utilized to analyze, model and predict response of 

trees to climate change. Although seldom appreciated, provenance research might be among the most important contributions of forestry to biological sciences.

561. Maschinski J, Ross MS, Liu H, O’Brien J, 

Wettberg EJ, Haskins KE. 2011. Sinking ships: conservation options for endemic taxa threatened by sea level rise. 

Climatic Change 107:147-167.

Type: Journal 

Geographic Area: Florida Keys, USA

Compilers’ Keywords: climate change, endangered species, 

extinction

Abstract: Low-elevation islands face threats from sea level rise 

(SLR) and increased storm intensity. Evidence of endangered 

species’ population declines and shifts in vegetation communities are already underway in the Florida Keys. SLR predictions indicate large areas of these habitats may be eliminated 

in the next century. Using the Florida Keys as a model system, 

we present a process for evaluating conservation options for 

rare and endemic taxa. Considering species characteristics and 

habitat, we assess central issues that influence conservation options. We contrast traditional and controversial options for two 

animal and two plant species giving special emphasis to perceptions of ecological risk and safety from SLR and suggest 

courses of action. Multiple strategies will be required to spread 

extinction risk and will be effective for different time periods. 

Global climate change presents an uncertain, perhaps no-analog future that will challenge land managers and practitioners 

to re-evaluate equilibrium-state-conceived laws and policies 

not only for these taxa, but for many facing similar threats. To 

embrace conservation in a changing world will require a new 

dialogue that includes controversial ideas, a review of existing 

laws and policies, and preparation for the oncoming change.

Link: http://softel.fiu.edu/projects/Peer%20Reviewed%20

Journal%20Articles/1307-Maschinski.pdf

562. Matyas C. 1994. Modeling climate change effects with 

provenance test data. Tree Physiology 17:797-804.

Type: Journal 

Geographic Area: North America

Keywords: adaptation, distribution, ecological distance, 

growth, species survival, successional change

Summary: Provenance tests of forest trees, which were originally intended to identify suitable seed sources for planting at 

different locations, provide valuable data for assessing the response of populations to environmental change. Environmental 

differences between the location of origin and the planting 

(test) site have been calculated by principal component analysis 

and termed ecological distance. Based on ecological distance 

values, the growth response of tree populations can be modeled as a function of the test site macroclimate. These models 

can then be used to predict the effects of climatic change on 

growth and survival. The growth response model predicts that 

increasing annual mean temperatures will result in accelerated 

growth if precipitation is sufficient, but only within the limits 

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 201

are: targeting species and habitats for seed collecting, facilitation of collecting and of reintroduction of species. The current 

limitations to the accessibility of this technology for seed conservation are discussed.

Link: http://www.kew.org/ucm/groups/public/documents/

document/ppcont_013766.pdf

566. O’Neill GA. 2013. Seed transfer 2.0: assisting assisted 

migration. In: Assisted Migration: A Primer for Reforestation 

and Restoration Decision Makers; Portland, OR. 51 p.

Type: Presentation

Geographic Area: North America

Compilers’ Keywords: Assisted Migration Adaptation Trial, 

climate change, forest management, seed transfer guidelines, 

seed transfer zones, genecology, provenance tests, seed transfer distance

Abstract: A growing body of provenance test data highlights the importance of seed source selection in reforestation and also suggests that climate change will render many 

tree populations substantially maladapted in the near future. 

Maladaptation may result in significant changes in health, 

productivity and distributions of foundation tree species, with 

attendant impacts on ecosystem function, carbon sequestration and the forest industry. Assisting the natural migration of 

seed sources to ensure forests are adapted to future climates is 

emerging as a key strategy to mitigate anticipated maladaptation in forest trees. These observations have placed new demands on seed source selection systems in forestry, while the 

availability of geographic information systems, fine scale climate models, a new era of provenance tests, and new analytical 

techniques have revolutionized opportunities for the development of new seed transfer systems. This presentation examines 

opportunities for creating new seed transfer systems and for 

implementing assisted migration, and discusses the design of 

seed transfer systems from the perspective of their structure, 

critical seed transfer distance calculation, and zone delineation 

methods, and ability to accommodate assisted migration. A 

hybrid fixed/focal point seed transfer system and methods to 

integrate assisted migration into existing fixed and focal point 

seed transfer systems will be proposed. 

Link: http://www.rngr.net/resources/assisted-migration

567. O’Neill GA, Ukrainetz NK, Carlson M, 

Cartwright C, Jaquish B, King J, Krakowski J, 

Russell JH, Stoehr MU, Xie CY, Yanchuk A. 2008.

Assisted migration to address climate change in British Columbia: 

recommendations for interim seed transfer standards. Victoria, 

British Columbia, Canada: B.C. Ministry of Forest and Range, 

Research Branch. 048. 38 p.

Type: Government Document

Geographic Area: Western Canada

564. McIntyre S. 2011. Ecological and anthropomorphic 

factors permitting low-risk assisted colonization in temperate 

grassy woodlands. Biological Conservation 144:1781-1789.

Type: Journal 

Geographic Area: Australia

Keywords: grasslands, eucalypt woodlands, plant conservation, translocation, managed relocation

Abstract: There is a risk that the potential of assisted colonization to contribute to biodiversity conservation will be lost 

in the storm of controversy that currently surrounds it. This 

paper describes a low-risk scenario for assisted colonization 

using plants. Using an analysis of temperature grassy woodlands from Australia, relevant ecosystem attributes are identified which make assisted colonization a sensible strategy, and 

that may characterize other favorable situations globally. The 

contributing elements include: a biota adapted to resource to 

resource conservationism, a naturally connected landscape 

with component species having wide distributions over a large 

climatic gradient, current land use unrelated to endogenous 

disturbance regimes resulting in extensive replacement and 

modification of the ecosystem overs its entire range. Intensive 

agriculture can create a highly-disturbed and nutrient-enriched 

landscape matrix, which effectively fragments the species 

assemblage. Relocation of plant species within and close to 

their range is not going to create an invasive situation in these 

landscapes. Candidates for assisted colonization are forbs 

and interstitial grasses that have persisted over much of their 

range, but which have declined within that range due to land 

use impacts. The suggested priority receiving sites would be 

those with a moderate level of past modifications, now being 

managed for conservation and with low nutrient status. The 

proposed use of assisted colonization is a conservative strategy 

that will build on current conservation practice and greatly improve the prospects for native plants where climate change is 

superimposed on a fragmented plant community. 

565. Moat J, Smith PP. 2003. Applications of geographical information systems in seed conservation. Chapter 4. 

In: Smith RD, Dickie JB, Linington SL, Pritchard HW, Probert 

RJ, editors. Seed Conservation: Turning Science into Practice. 

London, United Kingdom: Royal Botanic Gardens. p 79-88. 

Type: Book Section

Geographic Area: Global

Compilers’ Keywords: GIS, remote sensing, seed collection, 

reintroduction

Summary: The use of Geographic Information Systems (GIS) 

in seed conservation has enormous potential. This paper examines the use of spatial technology (both GIS and remote sensing) in the context of seed conservation planning. It explores 

three major areas to which GIS technology can be applied, 

with practical and visual examples. These three applications 

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202 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

paper is not intended as an endorsement of the AM approach; 

however, we feel there is considerable value in discussing 

implementation issues at this point in the AM debate.

569. Richardson DM, Hellmann JJ, McLachlan JS, 

Sax DF, Schwartz MW, Gonzalez P, Brennan EJ, 

Camacho A, Root TL, Sala OE, Schneider SH, Ashe 

DM, Clark JR, Early R, Etterson JR, Fielder ED, Gill 

JL, Minteer BA, Polasky S, Safford HD, Thompson 

AR, Vellend M. 2009. Multidimensional evaluation of 

managed relocation. Proceedings of the National Academy of 

Sciences USA 106:9721-9724.

Type: Journal 

Geographic Area: Global

Keywords: assisted migration, climate change, conservation 

biology, conservation strategy, sustainability science

Abstract: Managed relocation (MR) has rapidly emerged as 

a potential intervention strategy in the toolbox of biodiversity 

management under climate change. Previous authors have 

suggested that MR (also referred to as assisted colonization, 

assisted migration, or assisted translocation) could be a lastalternative option after interrogating a linear decision tree. 

We argue that numerous interacting and value-laden considerations demand a more inclusive strategy for evaluating MR. 

The pace of modern climate change demands decision making 

with imperfect information, and tools that elucidate this uncertainty and integrate scientific information and social values are 

urgently needed. We present a heuristic tool that incorporates 

both ecological and social criteria in a multidimensional decision-making framework. For visualization purposes, we collapse these criteria into 4 classes that can be depicted in graphical 2-D space. This framework offers a pragmatic approach for 

summarizing key dimensions of MR: capturing uncertainty in 

the evaluation criteria, creating transparency in the evaluation 

process, and recognizing the inherent tradeoffs that different 

stakeholders bring to evaluation of MR and its alternatives.

Link: http://www.ncbi.nlm.nih.gov/pubmed/19509337

570. Stapley J, Reger J, Feulner PG, Smadja 

C, Galindo J, Ekblom R, Bennison C, Ball AD, 

Beckerman AP, Slate J. 2010. Adaptation genomics: the 

next generation. Trends in Ecology and Evolution 25:705-712.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: animals, biological evolution, genomics, plants

Abstract: Understanding the genetics of how organisms adapt 

to changing environments is a fundamental topic in modern 

evolutionary ecology. The field is currently progressing rapidly 

because of advances in genomics technologies, especially DNA 

sequencing. The aim of this review is to first briefly summarize 

Compilers’ Keywords: seed transfer guideline, forest management, policies

Summary: Climate change is expected to result in trees 

in most regions of British Columbia becoming increasingly maladapted to the climates in which they are planted. 

Consequently, planting seedlings adapted to future climates 

(assisted migration) is recognized as a key strategy to address 

climate change, as it will help maintain healthy, productive 

forests, and ensure capture of gains obtained from decades of 

selective breeding. To examine opportunities to incorporate assisted migration into British Columbia’s seed transfer system, 

the feasibility of increasing the upper elevational transfer limit 

of British Columbia’s Class A and Class B seed was assessed 

by calculating the climatic transfer distance associated with elevational transfers. A rationale was developed for quantifying 

an appropriate climatic distance and range to migrate seed, and 

was used to evaluate elevational transfer increases of 100 and 

200 m. Results indicate that of the 30 Class A Seed Planning 

Units (SPUs) examined, eight should retain their current upper 

elevation limits, one should have their upper elevation limit 

increased by 100 m, and the remainder should have their upper 

elevation limits increased by 200 m for eight species, by 100 

m for two species, and should remain unchanged for three species. Specific recommendations are provided in Tables 2 and 3. 

Deployment of orchard seed in the lowest 200 m of the western white pine—Maritime and interior spruce—East Kootenay 

SPUs is discouraged, as is transfer of Class B seed of amabilis 

fir and western hemlock more than 200 m downward and western redcedar more than 300 m downward. 

Link: http://www.for.gov.bc.ca/hfd/pubs/Docs/Tr/Tr048.pdf

568. Pedlar J, McKenney DW, Beaulieu J, Colombo 

S, McLachlan JS, O’Neill GA. 2011. The implementation of assisted migration in Canadian forests. The Forestry 

Chronicle 87:766-777.

Type: Journal 

Geographic Area: Canada

Keywords: assisted migration, implementation, provenance 

data, seed procurement, Canada, forest, seed transfer, climate 

change

Abstract: We outline the major steps involved in implementing assisted migration (AM) and assess, in a general way, the 

capacity to carry out each step in Canadian forests. Our findings 

highlight the fact that capacity to implement AM differs between forest species; in particular, the existence of established 

provenance trials, seed transfer guidelines, seed procurement 

systems, and plantation establishment protocols makes AM 

considerably more feasible for most commercial tree species 

than for most species of conservation concern. We report on 

several AM efforts involving commercial tree species that are 

already underway in Canada and identify a number of initiatives that could be undertaken to help build AM capacity. This 

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 203

moving down out of Canada. Seeing the evidence of climate 

change prompts many conservation and restoration professionals to ask: what can we do to help our natural systems adapt 

to these rapid catastrophic changes? One common answer is 

assisted migration of taxa unable to move or adapt rapidly 

enough. But, how do we identify which taxa? Or where to 

move them? To undertake a successful assisted migration program requires the synthesis of multiple lines of evidence that 

cut across disciplines. We will present the most common tools 

that are currently being employed to understand the potential 

impacts of climate change on select taxa of plants, and provide 

the background necessary to evaluate their effectiveness. 

Link: http://www.rngr.net/resources/assisted-migration

573. Vitt P, Havens K, Hoegh-Guldberg O. 2009.

Assisted migration: part of an integrated conservation strategy. 

Trends in Ecology and Evolution 24:473-474.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: response letter, biological invasions, 

assisted colonization

Compilers’ Summary: A response letter to an article written 

by Ricciardi and Simberloff (2009). “Although we agree with 

several of their points, we believe that assisted migration could 

become an important part of integrated conservation strategies. Ricciardi and Simberloff focus on extreme examples 

of species translocations, including the ‘re-wilding’ of North 

America with African mammals and other examples of intercontinental and long distance transport of species. We do not 

condone these practices as legitimate conservation strategies 

because we also recognize the risk of biological invasions.”

Link: http://noss.cos.ucf.edu/papers/Vitt%20et%20al%20

2009%20and%20Schwartz%20et%20al%202009.pdf

574. Vitt P, Havens K, Kramer AT, Sollenberger D, 

Yates E. 2010. Assisted migration of plants: changes in latitudes, changes in attitudes. Biological Conservation 143:18-27.

Type: Journal 

Geographic Area: Great Lakes, USA, Global

Keywords: assisted migration, assisted colonization, managed relocation, climate change, range shifts, seed banking, 

biodiversity conservation

Abstract: Rapid climate change has the potential to alter the 

location of bioclimatic envelopes for a significant portion of the 

world’s flora. Plant species will respond variously via phenotypic plasticity, evolutionary adaptation, migration, or extinction. When fragmentation limits migration potential of many 

species or when natural migration rates are outstripped by the 

pace of climate change, some propose purposeful, human mediated migration (assisted migration) as a solution. Here, we 

join the debate on assisted migration, and while recognizing 

how next generation sequencing (NGS) has transformed our 

ability to identify the genes underpinning adaptation. We then 

demonstrate how the application of these genomic tools to ecological model species means that we can start addressing some 

of the questions that have puzzled ecological geneticists for 

decades such as: How many genes are involved in adaptation? 

What types of genetic variation are responsible for adaptation? 

Does adaptation utilize pre-existing genetic variation or does 

it require new mutations to arise following an environmental 

change?

Link: http://www.ncbi.nlm.nih.gov/pubmed/20952088

571. Ungerer MC, Johnson LC, Herman MA. 2008.

Ecological genomics: understanding gene and genome function 

in the natural environment. Heredity 100:178-183.

Type: Journal 

Geographic Area: Global

Keywords: ecological genomics, ecology, genomics, functional genomics, interdisciplinary, multidisciplinary

Abstract: The field of ecological genomics seeks to understand 

the genetic mechanisms underlying responses of organisms to 

their natural environments. This is being achieved through the 

application of functional genomic approaches to identify and 

characterize genes with ecological and evolutionary relevance. 

By its very nature, ecological genomics is an interdisciplinary 

field. In this review, we consider the significance of this new 

area of study from both an ecological and genomic perspective 

using examples from the recent literature. We submit that by 

considering more fully an ecological context, researchers may 

gain additional insights into the underlying genetic basis of 

ecologically relevant phenotypic variation. Likewise, genomic 

approaches are beginning to offer new insights into higher-level biological phenomena that previously occupied the realm of 

ecological investigation only. We discuss various approaches 

that are likely to be useful in ecological genomic studies and 

offer thoughts on where this field is headed in the future.

Link: http://www.ncbi.nlm.nih.gov/pubmed/17519970

572. Vitt P. 2013. Using multiple lines of evidence to prioritize assisted migration of both rare and common species. In: 

Assisted Migration: A Primer for Reforestation and Restoration 

Decision Makers; Portland, OR. 47 p.

Type: Presentation

Geographic Area: USA

Compilers’ Keywords: MaxEnt, species vulnerability, climate change, species mapping, management

Abstract: The weather just keeps getting more and more 

strange, record heat in 2012, coupled with storms and fires 

causing billions of dollars in damage. And 2013 began with 

freak tornados caused by unseasonably warm temperatures in 

the northern U.S. colliding with the more typical colder weather 

CHAPTER 3 • MIGRATION • STRATEGIES
204 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

may potentially offer a suite of complementary, precision tools 

for guiding seed source selection assisted migration or other 

reforestation needs. This talk outlines the scientific basis of 

landscape genomics, reviews results of early studies in the 

field and discusses potential applications. 

Link: http://www.rngr.net/resources/assisted-migration

576. Williams MI. 2013. Charting assisted migration as 

a climate change adaptation strategy. In: Assisted Migration: 

A Primer for Reforestation and Restoration Decision Makers; 

Portland, OR. 40 p.

Type: Presentation

Geographic Area: USA

Compilers’ Keywords: review, native plants, native plant 

transfer guidelines, seed transfer zones

Abstract: Assisted migration, defined as the movement of native plants to facilitate natural range expansion in direct management response to climate change, has gained attention as an 

adaptation strategy since 2007. It can occur as the movement 

of 1) seed sources and populations within their current range 

(assisted population migration), 2) seed sources and populations from their current range to suitable areas immediately 

adjacent (assisted range expansion), and 3) species to locations 

far outside their current range to curtail extinction (assisted 

species migration). Across federal, state, and private groups in 

the U.S., very few adaptation and mitigation plans have been 

implemented or evaluated despite the increase in the amount of 

management planning during the past 30 years. Uncertainties 

about future climate conditions, risks with moving species and 

populations outside their current ranges, and existing policies 

have hampered formal actions. Drawing from conventional 

reforestation practices and proposed adaptation strategies, we 

chart the implementation of assisted migration and provide resources for researchers, landowners, and nurseries to facilitate 

collaboration and development of reforestation and restoration 

plans.

Link: http://www.rngr.net/resources/assisted-migration

577. Williams MI, Dumroese RK. 2013. Preparing 

for climate change: forestry and assisted migration. Journal of 

Forestry 111:287-297.

Type: Journal 

Geographic Area: North America

Keywords: assisted migration, climate change, forest management, seed transfer, seed zones

Abstract: Although plants have moved across the landscape 

in response to changing climate for millennia, projections of 

contemporary climate change suggest that forest tree species 

and populations will need to migrate faster than their natural 

ability. Therefore, climate change adaptation strategies, such 

as assisted migration, have gained attention since 2007. 

the potential negative impacts, present a strategy to collect and 

bank seeds of plant species at risk of extinction in the face of 

rapid climate change to ensure that emerging habitats are as 

species-diverse as possible. We outline the framework currently being used by the Dixon National Tallgrass Prairie Seed 

Bank to prioritize species for seed banking, both for restoration 

purposes and for potential assisted migration in the future. We 

propose a strategy for collecting across the entirety of a species 

range, while targeting populations likely to go extinct under 

climate change, determined by application of species distribution models. Finally, we discuss current international efforts to 

collect and bank the global flora, as well as the research needs 

necessary to fully undertake the strategy presented. 

Link: http://www.researchgate.net/publication/222155402_

Assisted_migration_of_plants_Changes_in_latitudes_changes_in_attitudes/file/9c96051a4fcaab7382.pdf

575. Wheeler NC, Neale DB. 2013. Landscape genomics: genetic tools to inform seed source decisions for assisted migration. In: Assisted Migration: A Primer for Reforestation and 

Restoration Decision Makers; Portland, OR. 22 p.

Type: Presentation

Geographic Area: Global

Compilers’ Keywords: landscape management, genetic adaptation, climate change, tool

Abstract: Climate change in North America, characterized 

by warming trends and some shifts in precipitation patterns, is 

now a generally accepted reality. Evidence of shifting natural 

ranges, both expansion and contraction, can already be seen 

in forest trees. Assisted migration offers a means to maintain 

forest productivity and ecological function by expediting species movements. Identifying which seed sources to move to 

optimize goals will not be trivial task. Those managing plantation forests deal with few species and typically have extensive 

knowledge of their planting stock. Intensive tree improvement 

programs, based on extensive genetic evaluation trials over 

the landscape provide excellent guides to which seed sources 

may be moved, and how far, to optimize program goals. In 

recent years, traditional tree improvement approaches have 

been supplemented with tools that use a rapidly expanding 

genomic resource and database. Public land managers are 

typically not so fortunate in the genetic resources available to 

guide them in issues concerning reforestation following disturbance, habitat restoration, conservation or assisted migration. 

With respect to reforestation, decisions regarding seed source 

have typically relied on natural seeding in, seed zone collections, or in special cases, information gleamed from common 

garden or provenance trials. Neither of the first two methods 

is guaranteed to produce a standing crop adapted to the future 

environment. Good provenance trials can significantly improve those chances, but offer a number of fiscal and logistical 

hurdles. Genomic studies, conducted at the landscape level, 

CHAPTER 3 • MIGRATION • STRATEGIES
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 205

580. Winder R, Nelson EA, Beardmore T. 2011.

Ecological implications for assisted migration in Canadian forests. The Forestry Chronicle 87:731-744.

Type: Journal 

Geographic Area: Canada

Keywords: adaptation, assisted migration, climate change, 

colonization, constraints, ecology, establishment, mitigation, 

translocation, trees

Abstract: Forest ecosystems are already being impacted by 

climate change as natural migration rates are outpaced by 

rapidly changing climate conditions. Human assisted migration has been proposed as a potential management option to 

maintain optimal health and productivity of Canada’s forests; 

however, a better understanding of the ecological implications 

is needed to inform decision-making on assisted migration 

(AM). This paper examines the ecological constraints and 

consequences of AM, and discusses options for their mitigation at three scales: translocation over long distances (assisted 

long-distance migration), translocation just beyond the range 

limit (assisted range expansion), and translocation of genotypes within the existing range (assisted population migration). 

From an ecological perspective, we find that AM is a feasible 

management option for tree species and that constraints and 

consequences can be minimized through careful application of 

available knowledge and tools.

Effective implementation of assisted migration can only occur 

if target transfer guidelines are developed because our current 

seed transfer guidelines, established to guide the movement of 

plant materials, lack inherent spatial and temporal dynamics 

associated with climate change. This limitation restrains reforestation practitioners from making decisions about assisted 

migration. Lack of operating procedures, uncertainties about 

future climate conditions, risks associated with moving plants 

outside their current ranges, and existing policies have hampered formal actions in forest management and conservation. 

We review the current thinking on assisted migration of forest tree species and provide information that could facilitate 

implementation.

Link: http://www.treesearch.fs.fed.us/pubs/44260

578. Williams MI, Dumroese RK. 2013. Climate 

change and assisted migration: strategic options for forest and 

conservation nurseries. Forest Nursery Notes 33:33-35.

Type: Newsletter

Geographic Area: North America

Compilers’ Keywords: forest nurseries, management

Introduction: In light of current studies that show climate 

will change faster than plants can adapt or migrate naturally, 

it begs the question, “What does this mean for forestry, specifically forest and conservation nurseries?” In this article, we 

discuss options for forest nurseries, such as assisted migration. 

Link: http://www.treesearch.fs.fed.us/pubs/43883

579. Williams MI, Dumroese RK. 2014. Role of climate change in reforestation and nursery practices. Western 

Forester 59:11-13.

Type: Newsletter

Geographic Area: North America

Compilers’ Keywords: forest management, nursery practices, assisted migration, seed transfer, Society of American 

Foresters

Introduction (2nd paragraph): The divergence in rates 

between climate change and tree adaptation will have important consequences for reforestation and nursery practices. Plant 

materials outplanted today must be able to meet and face the 

climatic challenges during this century. Unfortunately, most 

state and commercial nurseries in the U.S. have not yet explored how changes in climate will impact their operations. 

This article highlights some adaptation strategies to help reforestation and nursery practices, such as moving populations to 

new locations, modifying seed transfer guidelines, and targeting diversity in plant materials.

Link: http://www.forestry.org/northwest/westernforester/2014/

CHAPTER 3 • MIGRATION • STRATEGIES
206 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

581. Center for Plant Conservation. Reintroduction 

registry. (URL accessed 29 November 2015) 

Type: Web Page

Geographic Area: USA

Compilers’ Keywords: conservation, extinction, plant species, online database, repository, restoration, Endangered 

Species Act

About: The Center for Plant Conservation (CPC) is dedicated 

solely to preventing the extinction of U.S. native plants. The 

Center was one of the first organizations created to meet this 

need. The Center is a network of 39 leading botanic institutions. 

Founded in 1984, the Center operates the only coordinated national program of off-site (ex situ) conservation of rare plant 

material. This conservation collection ensures that material is 

available for restoration and recovery efforts for these species. 

CPC also works in research, restoration, technical assistance, 

education and advocacy through the efforts of the network and 

the national office. The cooperative CPC network maintains 

the National Collection of Endangered Plants. Believed to be 

the largest living collection of rare plants in the world, the collection contains more than 750 of America’s most imperiled 

native plants. Live plant material is collected from nature under 

controlled conditions and then carefully maintained as seed, 

rooted cuttings or mature plants. Network institutions conduct 

horticultural research and carefully monitor these materials 

so that imperiled plants can be grown and returned to natural 

habitats. Several CPC institutions are also involved in restoration projects in the field (in situ). Scientists are stabilizing 

current populations of imperiled plants and reintroducing new 

populations in appropriate habitats. These conservation efforts 

are undertaken to complement other preservation activities for 

our nation’s flora, such as habitat protection and management. 

Off-site storage and cultivation of genetically appropriate plant 

material is a critical step in supporting restoration in the wild. 

The Center’s goal is to protect the most imperiled U.S. plants 

from extinction and restore them to secure habitats in cooperation with multiple conservation agencies and organizations. 

Almost 1,000 U.S. plant species are already listed under the 

federal Endangered Species Act or qualify for listing. Without 

human intervention, many of these species may be gone within 

our lifetime. Research by CPC has shown that 80 percent of the 

at-risk plants of the United States are closely related to plants 

with economic value somewhere in the world, and more than 

50 percent are related to crop species.

Link: http://www.centerforplantconservation.org/welcome.

asp

General (339–380)—debates, reviews, 

trends, surveys, summaries

Research (381–540)—migration studies, 

range shifts, reintroductions

Strategies (541–580)—adaptation options, 

frameworks, decision support

Resources (581–582)—tools, websites, 

software

Migration

#. Author(s), Date. Title. Source.

Type: 

Geographic Area: 

Keywords: 

Description:

Link:

location of study

keywords written by the 

author(s) or provided by the 

compilers

ID number and

information

abstract, summary, or 

portions of the introduction 

or conclusion written by the 

author(s) or provided by the 

compilers

book, book section, conference 

paper, dissertation, government 

document, journal article, 

newsletter, presentation, report, 

thesis or web page

internet address of reference

Each reference contains the following:

CHAPTER 3 • MIGRATION • RESOURCES
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 207

582. Maschinski J, Albrecht MA, Monks L, Haskins 

KE. 2012. Center for plant conservation best reintroduction 

practice guidelines. Appendix 1. In: Maschinski J, Haskins 

KE, editors. Plant Reintroductions in a Changing Climate: 

Promises and Perils. Washington (DC): Island Press. p 30. 

Type: Book Section

Geographic Area: Global

Compilers’ Keywords: climate change, plant restoration, 

translocation

Introduction (2nd paragraph): This is a quick reference 

for practitioners to use when planning and executing rare plant 

reintroductions. The term reintroduction in this appendix implies any attempt to introduce propagules to an unoccupied 

patch, including augmentations, introductions, and translocations. Managed relocations would require following these 

same guidelines in addition to the points presented by Haskins 

and Keel (this volume) and the modeling, interdisciplinary, 

multiagency, and potentially international collaborations cautioned by Kennedy and colleagues (this volume). The sections 

are intended to help practitioners do the following: justify the 

decision to conduct a reintroduction, prepare the reintroduction 

design with legal, funding, species biology, horticulture, and 

recipient site considerations in mind, implement the reintroduction, conduct project aftercare, and design monitoring to 

document long-term establishment of the rare population. All 

phases of the reintroduction process should include opportunities for public involvement. In addition, we suggest a template 

to use for documenting all aspects of the reintroduction that 

can be found on the North Carolina Botanical Garden website.

CHAPTER 3 • MIGRATION • RESOURCES
208 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

583. Auld GS, Stead JW, Aitken SN, Bull GQ. 2002.

Forest genetics and certification: global and local issues facing 

tree breeders, policy-makers, and forest managers in British 

Columbia. Vancouver, British Columbia, Canada: Centre for 

Forest Gene Conservation, University of British Columbia. 

200 p.

Type: Report

Geographic Area: Western Canada

Compilers’ Keywords: forest management, seed transfer

Executive Summary: Since the early 1990s, forest certification has become an internationally recognized mechanism for 

assessing and encouraging the practice of responsible forest 

management. Markets are increasingly demanding independent assurances that forest products come from sustainable 

sources. British Columbia (BC) forest managers, the provincial government and other stakeholders are keenly aware of 

these pressures; creating significant interest in assessing how 

BC’s present forest management regulations, standards, requirements and practices compare to the requirements set by 

the various certification initiatives. In this report, we examine 

one such issue. Our attention focuses on the way four different forest certification initiatives address the conservation and 

management of forest genetic resources. The intention being 

two-fold: 1) Describe how certification has developed internationally to deal with forest genetics issues and; 2) Relate 

these findings to the BC situation to uncover the current and 

potential future issues that face BC tree breeders, silviculturalists, forest companies, forest managers, and policy makers as 

they attempt to evaluate the application of certification to how 

BC’s forests are managed.

584. Curtis RO, DeBell DS, Miller RE, Newton M, 

St. Clair JB, Stein WI. 2007. Silvicultural research and the 

evolution of forest practices in the Douglas-fir region. Portland 

(OR): USDA Forest Service, Pacific Northwest Research 

Station. General Technical Report PNW-GTR-696. 172 p.

Type: Government Document

Geographic Area: Pacific Northwest USA

Keywords: forest history, silviculture, Douglas-fir, forest research, Pseudotsuga menziesii

Abstract: Silvicultural practices in the Douglas-fir region 

evolved through a combination of formal research, observation, and practical experience of forest managers and silviculturists, and changing economic and social factors. This process 

began more than a century ago and still continues. It has had a 

General (583–616)—reviews, history, 

summaries, justification

Research (617–802)—genetic studies, tree 

improvement, climate change

Strategies (803–827)—framework, delineation, modelling

Resources (828–877)—guidelines, zones, 

maps, tools, software

Transfer Guidelines

and Zones

CHAPTER 4 • TRANSFER GUIDELINES AND ZONES • GENERAL

#. Author(s), Date. Title. Source.

Type: 

Geographic Area: 

Keywords: 

Description:

Link:

location of study

keywords written by the 

author(s) or provided by the 

compilers

ID number and

information

abstract, summary, or 

portions of the introduction 

or conclusion written by the 

author(s) or provided by the 

compilers

book, book section, conference 

paper, dissertation, government 

document, journal article, 

newsletter, presentation, report, 

thesis or web page

internet address of reference

Each reference contains the following:
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 209

Link: http://www.treesearch.fs.fed.us/pubs/6939

586. Friedman ST, Foster GS. 1997. Forest genetics 

on federal lands in the United States: public concerns and policy 

responses. Canadian Journal of Forest Research 27:401-408.

Type: Journal 

Geographic Area: USA

Compilers’ Keywords: tree improvement, public land, public concern

Abstract: The challenge of managing public lands is to balance the current and future needs of people and the long-term 

health of ecosystems. In addition, public land management 

requires being responsive to the public and their elected representatives, and ensuring that public agencies are efficient, 

effective, and representative of the diversity of the American 

public. In this paper, the results of an informal survey of key 

stakeholders, and U.S. Forest Service employees with direct 

public contact are summarized. There are a host of public concerns about genetic diversity and tree improvement on public 

land. At the broadest level, species choice and the decision to 

artificially or naturally regenerate are issues in their own right, 

and yet are often linked to genetic concerns. If artificial regeneration is an option, there is the general concern about the 

impacts of changes in the genetics of future trees on components of the ecosystem. These concerns lie in three main areas: 

(1) reduction of genetic diversity, (2) loss of adaptation, and 

(3) changes in other ecosystem components from the directed 

selection of trees. At the current time, citizens range widely in 

their knowledge and concern about these issues. Developing 

policy to address public concerns can be approached through 

a variety of perspectives. Silvicultural techniques, including 

genetic resource management, are tools toward broader policy 

objectives for public land management. These objectives are 

currently the subject of much debate and controversy. Given 

that the broader policy context is dynamic, some policy options for genetics in public land management include (1) gene 

conservation only, (2) no frills, (3) appropriate technology with 

public support, and (4) application of economic criteria. Also 

discussed is the need for a framework to coordinate local and 

national policies regarding gene resource management.

587. Isaac LA. 1952. Advantages of selecting tree seeds 

with care. Tree Planters’ Notes 11:1-8.

Type: Government Document

Geographic Area: USA, Canada, Europe

Compilers’ Keywords: seed selection, forest management, 

reforestation, forest genetics, nursery stock

Introduction: The vast virgin forests of North America 

are gradually being cut away and converted into managed 

stands. The movement is well under way in the United States 

and advancing in the same direction in Canada. For the most 

great influence on the economic well-being of the region and 

on the present characteristics of the region’s forests. This long 

history is unknown to most of the public, and much of it is 

unfamiliar to many natural resource specialists outside (and 

even within) the field of silviculture. We trace the history of 

how we got where we are today and the contribution of silvicultural research to the evolution of forest practices. We give 

special attention to the large body of information developed in 

the first half of the past century that is becoming increasingly 

unfamiliar to both operational foresters and—perhaps more 

importantly—to those engaged in forestry research. We also 

discuss some current trends in silviculture and silviculturerelated research.

Link: http://www.treesearch.fs.fed.us/pubs/27615 

585. Everett RL. 1994. Restoration of stressed sites, 

and processes. Portland (OR): USDA Forest Service, Pacific 

Northwest Region. General Technical Report PNW-GTR-330. 

Volume IV. 123 p.

Type: Government Document

Geographic Area: Pacific Northwest USA

Keywords: restoration, forest health, ecosystem processes, 

disturbance effects, ecosystem management, insects, disease, 

and fire hazard

Abstract: Portions of forest ecosystems in eastern Oregon 

and Washington are in poor health, are not meeting societies’ 

expectations, and have elevated hazard for fire, insects, and 

disease. Diversity in stream habitats and associated fisheries 

has declined over the last several decades in several drainage 

basins, requiring conservation and restoration efforts in key 

watersheds. Required first steps in restoring forest and aquatic 

ecosystems are the immediate reduction in hazard for catastrophic loss of biodiversity, site quality, resource commodities, and improved conditions for public health. To prevent loss 

of future options we need to simultaneously reestablish ecosystem processes and disturbance effects that create and maintain desired sustainable ecosystems, while conserving genetic, 

species, community, and landscape diversity and long-term 

site productivity. Restoration of stressed sites is site specific, 

but the context for the action should be defined by the desired 

condition(s) of the next higher landscape scale and achieve 

desired positive cumulative effects over time. Restoration actions should be consistent with the desired level of disturbance 

effects required to maintain sustainable ecosystems, and standards and guides should reflect the inherent variability associated with dynamic systems. Costs associated with restoration 

activities should be weighed against the foregone benefits if no 

action is taken. The restoration of the biological components 

of ecosystems should provide increased opportunities for the 

restoration of human cultural, social, and economic ecosystem components and increase options for resource-dependent 

communities.

CHAPTER 4 • TRANSFER GUIDELINES AND ZONES • GENERAL
210 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

590. Johnson R, Stritch L, Olwell P, Lambert S, 

Horning ME, Cronn RC. 2010. What are the best seed 

sources for ecosystem restoration on BLM and USFS lands?

Native Plants Journal 11:117-131.

Type: Journal 

Geographic Area: USA

Keywords: seed zones, ecotypes

Abstract: Native plant restoration policy calls for use of “genetically appropriate” native plant material on USDI Bureau of 

Land Management (BLM) and USDA Forest Service (USFS) 

lands. In this article, we summarize experimental evidence 

showing that local adaptation is widespread in all kingdoms of 

life, and how this “home-field advantage” has been exploited 

in forest restoration activities to develop and promote regionally adapted, genetically diverse restoration materials. The 

importance of such materials is highlighted in the context of 

changing environmental conditions, such as those predicted 

in future climate change scenarios. Although the adaptive 

properties of most restoration species remains unexamined, 

numerous tools exist for identifying similar environments and 

ecosystems, and these can be used to develop first-generation 

seed zones. Finally, general recommendations for establishing 

foundation populations are outlined so that pre-adapted populations retain sufficient genetic diversity to acclimate to new 

evolutionary challenges, while minimizing the likelihood of 

deleterious genetic outcomes (for example, inadvertent selection of undesirable traits; inbreeding depression).

Link: http://www.treesearch.fs.fed.us/pubs/37836

591. Jones TA. 2005. Genetic principles and the use of native seeds—just the FAQs, please, just the FAQs. Native Plants 

Journal 6:14-24.

Type: Journal 

Geographic Area: Global

Keywords: genetics, adaptation, polyploidy, inbreeding 

depression, outbreeding depression, cross-pollination, selfpollination, apomixis, ecoregion, ecotype, seed transfer zones, 

cultivar, hybridization, selection

Abstract: To make intelligent choices in the marketplace, 

native seed customers should have a working understanding 

of genetic principles and terminology as they apply to selfpollinated, cross-pollinated, and apomictic plant materials. 

Customers should understand the genetic implications of a species’ breeding system, the various approaches used to decide 

what should be planted where, the risk of inbreeding or outbreeding depression, the meaning of commonly misunderstood 

terms such as “ecotype” and “cultivar,” and the role of hybridization and artificial selection in plant materials development. 

Plant material selection involves consideration of geographic 

(such as ecoregion, precipitation, winter hardiness, soil type), 

genetic (molecular markers), and adaptation (field testing) data.

part, foresters and timberland owners have been content to 

accept such regeneration as came in naturally regardless of 

parentage or even of species. For early artificial reforestation, any seed source was acceptable. But the rising costs of 

timber growing and increasing competition for land use have 

forced owners to recognize that it is necessary and profitable 

to secure the best seed for a given site. We discuss the strains 

of forest trees, climatic limitation of forest trees, climatic 

guides for seed selection, selection of stands and nursery 

stock, and finally provide steps for better seed. 

Link: http://www.rngr.net/publications/tpn/3-2/PDF.2004-10-

08.3621

588. Johnson R. 2000. Tree improvement in the Pacific 

Northwest. In: Rose R, Hasse DL. Proceedings: Advances and 

Challenges in Forest Regeneration. Corvallis (OR): Oregon 

State University. p 29-34.

Type: Book Section

Geographic Area: Pacific Northwest USA

Keywords: tree breeding, genetic gain, genetic diversity, disease resistance

Abstract: Advanced-generation tree breeding programs 

are underway for Douglas-fir and coastal western hemlock. 

These programs will continue to improve rates and other 

traits. Regardless of whether seed is from a seed orchard or 

natural collection, it must be used in its appropriate breeding 

zone or seed zone. These zones vary by species. Breeding 

programs are underway for other species as well, with many 

of these programs emphasizing disease and insect resistance. 

Absolute gains at rotation are still unknown, but absolute 

(not percentage) gains observed early in the rotation should 

increase to some degree with time. 

Link: http://www.treesearch.fs.fed.us/pubs/4921

589. Johnson GR, Sorensen FC, St. Clair JB, Cronn 

RC. 2004. Pacific northwest forest tree seed zones: a template 

for native plants? Native Plants Journal 5:131-140.

Type: Journal 

Geographic Area: Pacific Northwest USA

Keywords: seed movement, genetic variation, adaptation

Abstract: Seed movement guidelines for restoration activities are lacking for most native grasses, forbs, and shrubs. 

The forestry community has decades of experience in establishing seed zones and seed movement guidelines that may 

be of value to restoration managers. We review the history of 

seed zone development in forest trees, with emphasis on the 

Pacific Northwest, and make some suggestions concerning 

seed transfer guidelines for other native plants.

Link: http://www.treesearch.fs.fed.us/pubs/25517

CHAPTER 4 • TRANSFER GUIDELINES AND ZONES • GENERAL
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 211

Link: http://www.fs.fed.us/psw/publications/knapp/

psw_1994_knapp001.pdf

594. Knight TM, Havens K, Vitt P. 2011. Will the 

use of less fecund cultivars reduce the invasiveness of perennial 

plants? BioScience 61:816-822.

Type: Journal 

Geographic Area: Global

Keywords: invasive plant, ornamental plant, cultivar, 

demography

Abstract: Many invasive species were originally introduced 

for horticultural purposes, and several continue to be profitable for the green (nursery, horticulture, and landscape) industry. Recently, some plant suppliers have marketed less 

fecund cultivars of several invasive species, including glossy 

buckthorn (Frangula alnus), burning bush (Euonymus alatus), 

and Japanese barberry (Berberis thunbergii), as “safe” alternatives to invasive relatives. We use published matrix population models to simulate the effect of reducing fecundity on 

the population growth rates of invasive species. We show that 

large changes in fecundity result in relatively small changes to 

the population growth rates of long-lived species, which suggests that less fecund cultivars may still provide an invasive 

threat. Furthermore, many cultivars are clonal selections, and 

if crossed with other cultivars or selfed, they produce offspring 

with traits and fecundities that do not resemble the parent plant. 

On the basis of these two lines of evidence, we suggest that 

only female sterile cultivars that cannot reproduce asexually 

should be considered “safe” and noninvasive. Marketing less 

fecund cultivars as “safe” is premature at this time, and further 

research is necessary to determine the potential invasiveness of 

different cultivars.

Link: http://wubio.wustl.edu/files/biology/imce/knight_et_

al_2011_bioscience.pdf

595. Kramer AT, Havens K. 2009. Plant conservation genetics in a changing world. Trends in Plant Science 14:599-607.

Type: Journal 

Geographic Area: USA

Compilers’ Keywords: adaptation, biodiversity, endangered 

species, molecular evolution, genetic variation, population genetics, population dynamics, gene flow, seed transfer zones, 

species specificity, restoration, review, climate change

Abstract: Plant conservation genetics provides tools to guide 

conservation and restoration efforts, measure and monitor 

success, and ultimately minimize extinction risk by conserving species as dynamic entities capable of evolving in the face 

of changing conditions. We consider the application of these 

tools to rare and common species alike, as ongoing threats 

that increasingly limit their resilience, evolutionary potential 

and survival. Whereas neutral marker studies have contributed 

592. Kaye TN. 2001. Common ground and controversy 

in native plant restoration: the SOMS debate, source distance, 

plant selections, and a restoration-oriented definition of native.

In: Haase DL, Rose R, editors. Native Plant Propagation and 

Restoration Strategies. Portland (OR): Nursery Technology 

Cooperative and Western Forestry and Conservation 

Association. p 5-12.

Type: Conference Paper

Geographic Area: Global

Compilers’ Keywords: transfer

Abstract: Propagation and planting of native plants for 

habitat restoration is a multi-faceted process. There are 

many issues over which there is general agreement among 

restorationists, but there are a number of subjects that cause 

disagreement. For example, restorationists often agree that 

native plants should be emphasized, but disagree over where 

seeds or transplants should come from. In this paper, I examine four areas of controversy: the use single or multiple 

sources of a species at a given restoration site (the SOMS 

debate), source distance of plant materials, the use of native 

plant selections, and the importance of one’s definition of 

“native plant.” I conclude that some of these issues may be 

resolved through careful research, while others will remain a 

matter of personal opinion, and can only be resolved through 

a clear statement and scope of objectives of each restoration 

project.

Link: http://appliedeco.org/reports/restoration-source-genetics.pdf

593. Knapp EE, Rice KJ. 1994. Starting from seed: genetic issues in using native grasses for restoration. Restoration 

and Management Notes 12:40-45.

Type: Journal 

Geographic Area: USA

Compilers’ Keywords: genetic resources, seed collection, 

seed storage

Introduction: Historical records of the botanical composition 

of grasslands in their pristine state are often incomplete or lacking, yet it is evident that many native species were at one time 

much more widespread than they are today. Changing land use, 

including agricultural conversion, urbanization, overgrazing, 

fire suppression, and introduction of non-native weedy annuals, have all dramatically altered the landscape and the flora. In 

recent years, the public has become better informed about the 

decline of our natural heritage due to human influence, and the 

importance of biodiversity is not widely understood. Together 

with and perhaps in part due to this increasing public awareness has been a mounting interest in restoration using native 

species, including grasses. This paper will discuss, in a broader 

sense, some genetic issues that may be useful to consider when 

collecting and utilizing native grasses for restoration.

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second-hand quotation as far as possible and, what is still more 

important, not to accept other peoples’ labelling, criticism—

or laudation—of antecedent as well as contemporary authors 

uncritically. All of them have generally published much more 

material worthy of consideration than that which is usually attached to their names. The field is wide enough for bestowing 

on classical and other early authors the reference they by their 

works and writings are rightfully entitled to, as well as quoting 

contemporaries as they merit consideration. The old authors 

did not have the plenty of facts and experimental evidence as 

we have, and, above all, they were not strained by a publication explosion such as we now experience. They had, and took, 

more time to reflect and speculate upon the subjects within 

range of their knowledge. And one thing is evident: they were 

not a bit less intelligent that the authors of today. Darwin is an 

amazingly modern author, and he is in no way unique in this 

respect. Ecological experimental work is often laborious and 

frequently it does not elucidate the problems as perfectly as we 

hoped for. There is then nothing else to do than to proceed with 

new experiments and to follow the wisdom by the classical 

author, the famous Lamarck (in his Philosophie zoologique II, 

p. 174, 1809), here in the good translation by Greene (1959, p. 

158): “Let us collect with care the facts we can observe, let us 

consult experience wherever we can, and when this experience 

is inaccessible to us, let us assemble all the inductions which 

observation of facts analogous to those which escape us can 

furnisbh and let us assert nothing categorically; in this way, 

we shall be able little by little to discover the causes of a multitude of natural phenomena, and, perhaps, even of phenomena 

which seem the most incomprehensible...”

597. Mahalovich MF. 1997. Communicating the role 

of genetics in management. In: Communicating the Role of 

Silviculture in Managing the National Forests: Proceedings 

of the National Silviculture Workshop. Radnor (PA): USDA 

Forest Service, Northeastern Forest Experiment Station. 

General Technical Report NE-238. p 58-64.

Type: Government Document

Geographic Area: Global

Compilers’ Keywords: forest management, tree improvement, silviculture

Abstract: Three current issues serve as examples to convey the 

role of genetics in management. (1) Consequences of (qualitative) data. Results of simulated data for diameter (quantitative 

data) over several generations, illustrate the pitfalls of working 

towards balanced uneven-aged silvicultural systems in northern red oak, under natural regeneration constraints and existing 

management direction. (2) Comparisons of section boundaries 

within an ecological classification system and climatic zones 

(homoclines) as surrogates for managing genetic resources are 

of limited utility in describing patterns of genetic variation for 

adaptive, growth, and disease resistance traits. (3) Reporting 

gains and seed yields from tree improvement programs in 

much to conservation genetics, we argue for a renewed focus 

on quantitative genetic studies to determine how, or if, species 

will adapt to changing conditions. Because restoration plays an 

increasingly vital role in conservation, we discuss additional 

genetic considerations and research questions that must be actively studied now to effectively inform future actions.

Link: http://www.ncbi.nlm.nih.gov/pubmed/19748300

596. Langlet O. 1971. Two hundred years genecology. 

Taxon 20:653-721.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: genetic research, review, provenance 

trial, forest management, genotype, environmental interactions, climate change, local adaptation

Epilogue: This essay is a review of the research and literature 

concerning intraspecific diversity in ecologically significant 

features. In this category may be most of the relative importance and possibly also the course of influence of antecedent 

work. In a way it is tempting simply to summarize the result 

with the verse in the Ecclesiastes (I: 8): “... and there is nothing new under the sun.” But, indeed, there is. Roughly, there 

may be discerned three main stages of genecological work. 

First, the observation and statement of the mere existence of 

an intraspecific diversity in ecologically significant features. 

In this category may be most of the provenance work in respect to forest trees, agricultural and horticultural plants, as 

well as much of the investigations on herbaceous ecotypes, 

as Turrill (1940) put it: “The deservedly well-known experimental researches of Turesson must, for the most part as at 

present published, be considered under the heading of simple 

cultivation.” Second, the correlation of the diversity in ecologically significant features to the external environmental factors 

allows a preliminary estimation of the ecological importance 

of the extrinsic influences. Also, the patterns of intra-specific 

diversity are related to the patterns of environmental factors. 

The local interbreeding community or deme is found to be the 

basic genecological concept. Third, the intricate mechanisms 

of adaptations and the development of adaptations need to be 

experimentally investigated. Phytotrone or temperature-controlled growth-chamber experiments may be the next method 

to investigate the reactions of variously adapted populations 

to different external factors. Such work was first begun in the 

1950’s (cf. Hiesey 1953; Mooney & Billings 1961) and there 

is still much to be done and much to be gained in this field. 

Even if it may be of importance to delimit the ranges of different branches of biology from one another, as stressed above, it 

is equally important not to isolate these branches from teamwork. The great and real progress may be expected from the 

joint efforts of various specialists, not least from the cooperation of statisticians and computer specialists. I have learned one 

really important lesson in preparing this essay, viz., to avoid 

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The consequences of introducing stock without consideration 

of genetic variability are 1) the stock may die after planting 

because they lack adaptive characteristics, 2) delayed death; 

plants that evolved within the system already have the toolbox 

of adaptive genes, 3) poor growth, 4) inbreeding depression, 

and 5) genetic contamination. To curb these consequences, 

they recommend the following guidelines: 1) don’t buy planting stock unless there is no other option; instead collect it on 

or near the plant site but consider site conditions (elevation, 

slope, aspect, soil, drainages, frost dates, etc.), 2) collect from 

adjacent lands of similar topography and vegetation and be 

aware of changes in conditions that would favor genetic differences, 3) gather all information about the species’ genetic 

variation; when lacking match the collection sites with restoration sites by picking sites similar in elevation, soils, vegetation, 

and ecology, 4) collect from many parents at least 100 m apart 

to avoid inbreeding depression, 5) find a nursery that maintains 

records of stock origin, number of parent plants; if stock and 

origin unknown keep the numbers high and buy from several 

different nurseries, 6) give nature a chance first; sometimes it’s 

best not to intervene, and 7) if no material is available, build 

a new landrace. In any restoration effort, it is important to record location of planting site, origin, age and date of planting, 

survival and performance over time, and ecological behavior 

of stock.

Link: http://er.uwpress.org/content/7/1/18.full.pdf

600. Pike C. 2008. What is local? An introduction to genetics and plant selection in the urban context. Urban Habitats 

5:3-6.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: urban landscapes, inbreeding depression, assisted migration, local adaptation

Introduction: The urban landscape comprises myriad isolated green spaces inhabited by an assortment of vegetation 

types. To many city dwellers, these green spaces interrupt the 

monotony of concrete and steel and foster deep social attachment between city dwellers and nature. To a conservationist, 

these vegetation islands provide unique opportunities to restore ecological function to degraded areas by revegetating 

them with native plants. However, the restoration ecologist 

faces many challenges unique to the urban landscape. A significant investment in site preparation may be needed to offset the impacts of abiotic factors including compacted soil, 

drought, and air pollution. Seedlings that survive to maturity 

are not guaranteed immunity from these abiotic stresses, as is 

evident in the tree dieback and declines that plague many city 

landscapes. Abiotic factors are rarely the sole causal agent of 

urban tree declines since the presence of a multitude of other 

factors, such as insects and disease, are associated with symptoms of this decline. For example, drought in combination with 

Forest Service decision documents is recommended as means 

of showing consequences of ‘action vs. no action’ for genetic 

resources, thereby placing genetics in a more active role in the 

Agency’s next round of forest planning.

Link: http://www.treesearch.fs.fed.us/pubs/14994

598. McKay JK, Christian CE, Harrison S, Rice KJ. 

2005. “How Local Is Local?” A review of practical and conceptual issues in genetics of restoration. Restoration Ecology 

13:432-440.

Type: Journal 

Geographic Area: Global

Keywords: ecotype, local adaptation, outbreeding depression, 

restoration, seed zones

Abstract: In plant conservation, restoration (the augmentation or reestablishment of an extinct population or community) 

is a valuable tool to mitigate the loss of habitat. However, restoration efforts can result in the introduction of novel genes 

and genotypes into populations when plant materials used are 

not of local origin. This movement is potentially important because many plant species are subdivided into populations that 

are adapted to local environmental conditions. Here we focus 

on genetic concerns arising from ongoing restoration efforts, 

where often little is known about ‘‘How local is local?’’ (i.e., 

the geographic or environmental scale over which plant species are adapted). We review the major issues regarding gene 

flow and local adaptation in the restoration of natural plant 

populations. Finally, we offer some practical, common-sense 

guidelines for the consideration of genetic structure when restoring natural plant populations.

Link: http://www.des.ucdavis.edu/faculty/Harrison/people/

sue/papers/mckay_etal2005_RE.pdf

599. Millar CI, Libby WJ. 1989. Disneyland or native 

ecosystem: genetics and the restorationist. Restoration and 

Management Notes 7:18-24.

Type: Journal 

Geographic Area: USA

Compilers’ Keywords: seed transfer guidelines, forestry, genetic variation, cline, climate, seed zone, review, reforestation

Compilers’ Summary: Drawing from literature and their 

professional experience, the authors discuss the impacts of introduced genetic material in restoration plantings and propose 

general guidelines to avoid short and long term failures. Since 

the observation was made that seeds of different geographic origins produce trees with different survival and growth characteristics within the same plantation (HL Duhamel de Monceau 

of France in early 1700s), research forestry has supported two 

points: 1) species are genetically structured in a hierarchical 

fashion, and 2) variation can reflect adaptations to environmental conditions in which the ancestral populations evolved. 

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to our understanding of genetic variation in selected revegetation species within their range in Region One. The results of 

the study will be used to develop seed transfer guidelines for 

Forest Service System lands in Region One as well as adjacent 

areas. Results will also be used to compare native genotypes to 

“restoration cultivars.”

Link: http://www.fs.fed.us/wildflowers/nativeplantmaterials/

documents/npmreports/fy2008/R1/Seed_Transfer_Zone_

Common_Garden_Study.pdf

603. Rinehart S, Mahalovich MF. 2009. Development 

of seed transfer zones for Idaho fescue and bluebunch wheatgrass through common garden studies in region one. Missoula 

(MT): USDA Forest Service, Northern Region. 1 p.

Type: Government Document

Geographic Area: Northwestern USA

Compilers’ Keywords: native grass

Introduction: In 2008 Region One initiated a multi-year 

project to develop seed transfer zones for core native revegetation species. The project utilizes common garden study methodology and will contribute to our understanding of genetic 

variation in selected revegetation species within their range 

in Region One. The results of the study will be used to develop seed transfer guidelines for Forest Service System lands 

in Region One as well as adjacent areas. Results will also be 

used to compare native genotypes to “restoration cultivars.” 

During the 2008 field season, R1 forests and grasslands conducted wildland seed collection for two revegetation species, 

Idaho fescue (Festuca idahoensis) and bluebunch wheatgrass 

(Pseudoroegneria spicata).

Link: http://www.fs.fed.us/wildflowers/nativeplantmaterials/

documents/npmreports/fy2009/R1/Seed_Transfer_Zone_

Study_FEID_PSSP6.pdf

604. Rinehart S, Mahalovich MF. 2009. Development 

of seed transfer zones for two native shrubs through common 

garden studies in Region One. Missoula (MT): USDA Forest 

Service, Northern Region. 1 p.

Type: Government Document

Geographic Area: Northwestern USA

Compilers’ Keywords: Red osier dogwood, Shinyleaf spirea

Introduction: The Five-Year Action Plan for the R1 Native 

Plant Program identified the need to develop seed transfer 

zones for core native revegetation species. In 2008 a multiyear project was initiated to utilize common garden study 

methodology in collecting data on select native plant species 

commonly used for revegetation. This project will contribute 

to our understanding of genetic variation in native species 

within their range in the northern Rocky Mountain Ecosystem 

and northern Great Plains. The results of the study will be used 

to develop seed transfer guidelines for Forest Service System 

insect defoliation predisposed black oak (Quercus velutina) to 

decline, while defoliation and several pathogens were causal 

agents in the decline of English oak (Quercus robur). The impacts of biotic factors on plant health may be heightened or 

lessened in urban areas depending on the ecology of the insect 

or disease. In these examples, black oaks in urban areas were 

more susceptible to damage from gall wasps than trees surrounded by contiguous forest. In contrast, the pathogens inciting decline in Quercus robur are less problematic in urban 

settings where soil disturbance prevent Armillaria fungus from 

spreading great distances.

Link: http://www.urbanhabitats.org/v05n01/local_pdf.pdf

601. Powers RF. 1999. On the sustainable productivity of 

planted forests. New Forests 17:263-306.

Type: Journal 

Geographic Area: Global

Keywords: history of plantations, site potential, nutrition, soil 

properties, plantation yield

Abstract: Planted forests have more than a millennium of history and represent the world’s best hope for meeting global 

wood requirements in the twenty-first century. Advances in genetic improvement, nursery practices, stand establishment, and 

tending, harvesting, and manufacturing have boosted plantation yields to a higher level than at any point in history. Despite 

this, forest managers face a mounting challenge to demonstrate 

that plantation productivity is sustainable. Tackling this challenge requires a sound understanding of the principles of forest productivity, how they apply to a developing plantation, 

and how they are influenced by management. In this paper 

criticisms of plantation forestry are discussed from the basis 

of world experience, and examples of productivity decline are 

described. Obvious declines are rare, and can be attributed to 

poor soil management. However, ambiguities exist and controversy will continue until sustainable productivity can be 

demonstrated conclusively. Proposed programs aim to provide 

the technical base needed for sound soil management and sustainable plantation productivity.

Link: http://rothforestry.com/Resources/Powers1999.pdf

602. Rinehart S, Mahalovich MF. 2008. Development 

of seed transfer zones through common garden studies in Region 

One. Missoula (MT): USDA Forest Service, Northern Region. 

1 p.

Type: Government Document

Geographic Area: Northwestern USA

Compilers’ Keywords: native grass, Idaho fescue, bluebunch wheatgrass

Introduction: In 2008 Region One initiated a multi-year 

project to develop seed transfer zones for core native revegetation species. The R1 Common Garden Study will contribute 

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grasses. This shift to natives has raised questions about the 

selection of appropriate genetic stocks, similar to the issues 

involved in using native trees for reforestation. There is a need 

and an opportunity for dialog between land managers choosing 

native plant materials for wildland use, and geneticists who can 

provide counsel on relevant (but often complicated) details. 

This guide is intended to foster such a dialog. It synthesizes 

important genetic principles and provides many examples to 

better inform land managers, and help them understand the 

context for advice from geneticists. It focuses on details relevant to the USDA Forest Service’s Rocky Mountain Region, 

but contains a wealth of information that is relevant elsewhere, 

as well. Revegetation, restoration, or emergency rehabilitation 

efforts are conducted at the local level, but assistance can be 

provided from regional or national levels. Tools are needed to 

help local land managers make informed decisions when native materials suitable for the site are not commercially available (or feasible), and when seed transfer guidelines or seed 

zones have not been developed for the particular plant species 

of interest. Regional strategies can help provide guidance, and 

lead to economies of scale. Development of this Guide was 

the top priority in a strategic effort in the Rocky Mountain 

Region in 2002, to assist our National Forests and Grasslands 

in addressing their needs for native plant materials. The other 

products (for example, prioritized lists of species, seed procurement plans, native seed collections from the Forests and 

Grasslands, common garden studies) remain unfunded at this 

time. A tremendous amount of scientific study, policy, and infrastructure support the use of merchantable native tree species 

for reforestation following timber harvest, fire, disease or insect epidemics, and other disturbances. But we are in the early 

stages of developing and incorporating genetic information 

into our selection of other native plant materials in public land 

management. There is also a significant need to understand the 

interplay between our choices of germplasm and its ecological 

interaction in the landscape (for example, competitive interactions, pollination ecology, herbivory, etc.). It is hoped that 

this Guide will be further developed over time or lead to the 

production of other documents, workshops, policies or procedures, and serve as a catalyst to elevate the need (increase 

the awareness) for native seed collections and common garden 

studies.

Link: http://www.fs.fed.us/r2/publications/botany/plantgenetics.pdf

607. Saenz-Romero C, Snively AE, Lindig-Cisneros 

R. 2003. Conservation and restoration of pine forest genetic 

resources in Mexico. Silvae Genetica 52:233-237.

Type: Journal 

Geographic Area: Michoacán, Mexico

Keywords: Pinus, genetic resources, adaptive genetic variation, conservation, biodiversity, reforestation, ecological restoration, seed zones, Mexico 

lands in Region One as well as adjacent areas. Results will 

also be used to compare native genotypes to “restoration cultivars.” During the 2009 field season, Region One forests and 

grasslands conducted wildland seed collection for two shrub 

species: red osier dogwood (Cornus sericea ssp. sericea) and 

shinyleaf spirea (Spiraea betulifolia var. lucida). Common 

garden studies for these two species will be conducted by the 

Coeur d’Alene Nursery from 2010 through 2012. Seedlings 

will be grown out in the greenhouse in early spring, 2010, 

before being transferred to common garden study field plots. 

Data collection will involve plant morphology and phenology 

assessments at various plant growth stages.

Link: http://www.fs.fed.us/wildflowers/nativeplantmaterials/

documents/npmreports/fy2009/R1/Shrub_Seed_Transfer_

Zone_Study.pdf

605. Rogers DL. 2004. Genetic erosion: no longer just an 

agricultural issue. Native Plants Journal 5:113-122.

Type: Journal 

Geographic Area: Global

Keywords: genetic diversity, reforestation, restoration, revegetation, source materials, natural areas, conservation

Abstract: Genetic considerations pertaining to planting 

projects usually emphasize locally adapted source material. 

However, the amount of genetic diversity in the plant materials is important as well. Genetic erosion is the loss of genetic 

diversity—often magnified or accelerated by human activities. 

In native plant populations, genetic erosion results from habitat 

loss and fragmentation, but it also can result from a narrow genetic base in the original collections or by practices that reduce 

genetic diversity. Although species specific guidelines are not 

available, managers can minimize the risk of genetic erosion 

by being familiar with the biology of the affected species (including breeding system, mode of reproduction, and pattern of 

genetic diversity). Narrowly based genetic collections should 

be avoided, providers of plant materials for revegetation projects should offer information on their collection methods, 

and nursery managers should endeavor to minimize diversity 

losses at all stages of nursery culture.

Link: http://npj.uwpress.org/content/5/2/112.full.pdf+html

606. Rogers DL, Montalvo AM. 2004. Genetically appropriate choices for plant materials to maintain biological diversity. Davis (CA): University of California. 343 p.

Type: Report

Geographic Area: Northwestern USA

Compilers’ Keywords: genetic material, native plant material 

development, guidelines, seed zones, seed transfer guidelines

Forward: Native plant material use in revegetation or restoration efforts has increased nationally over the past couple of decades, replacing to some extent our past reliance on European 

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Maintaining genetic diversity in restoration materials is costly 

and does not have universal support. The use of provenance 

material (or local ecotypes) was hotly contested with strong 

and sound arguments on both sides of the issue. Procurement 

of pure ecotypes was difficult because of the variety of institutions involved in production and complications such as artificial selection and cross-pollination.

610. St. Clair JB, Johnson R. 2004. Structure of genetic variation and implications for the management of seed 

and planting stock. In: Riley LE, Dumroese RK, Landis TD, 

editors. Forest and Conservation Nursery Associations—2003. 

Fort Collins (CO): USDA Forest Service, Rocky Mountain 

Research Station. Proceedings RMRS-P-33. p 64-71.

Type: Government Document

Geographic Area: Western USA

Keywords: genetic structure, seed zones, breeding zones, 

seed transfer guidelines, genecology

Abstract: This paper reviews what is known about genetic 

structure of forest trees, and how that knowledge is used to 

determine safe limits to the movement of plant material. 

Geographic genetic variation in adaptive traits is of greatest 

importance to concerns of seed movement. Genetic structure 

in adaptive traits may be ascertained through long-term provenance and progeny tests, or short-term common garden studies 

in a nursery or nursery-like environment. These studies have 

shown that variation patterns are not consistent among species, among regions within a species, or among traits. The first 

seed zones were developed based on differences in climate and 

vegetation, and did not account for differences among species. 

Seed zones were recently revised in Oregon and Washington 

to reflect current knowledge of geographic genetic variation 

for individual species. Seed zones are an administrative convenience that directs managers how to bulk seeds from different 

stands. The use of seed transfer guidelines, on the other hand, 

allows greater flexibility and better knowledge of the risks of 

seed movement. Transfer guidelines, however, require keeping 

track of many small seed lots, which involves more time and 

expense.

Link: http://www.fs.fed.us/rm/pubs/rmrs_p033/rmrs_

p033_064_071.pdf

611. Turesson G. 1922. The species and the variety as ecological units. Hereditas 111:100-113.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: genomics, plant adaptation, 

biogeography

Introduction (4th paragraph): In the efforts made by the 

writer to arrive at an understanding of the Linnean species from 

an ecological point of view—of the ecospecies, as I prefer to 

Abstract: Deforestation rates in México are about 670,000 

ha/year. This threatens the richness of forest genetic resources 

in México, causing the disappearance of locally adapted populations and rare and endangered pine species. México is one 

of the six megadiverse countries in the world, with half of the 

world’s Pinus species. Pinus is one of the most economically 

and ecologically important forest genera in México. We suggest that delineation of seed zones and the establishment of 

a network of Forest Genetic Resource Conservation Units 

(FGRCUs), linked with forest management and ecological 

restoration programs will protect this valuable resource. We 

estimate that FGRCUs should include 25 to 50 ha each, with at 

least one FGRCU for each priority species in each seed zone. 

We highlight the need for studies of adaptive genetic variation among pine populations and for new methodologies and 

techniques to suit ecological restoration under Mexican forest 

conditions. We briefly describe ongoing research on these topics on forests owned by a well-organized indigenous community in Nuevo San Juan Parangaricutiro, Michoacán, western 

México.

608. Smith SE, Halbrook K. 2004. A plant genetics 

primer: basic terminology. Native Plants Journal 5:105-111.

Type: Journal 

Geographic Area: Global

Keywords: terminology, concepts, glossary

Abstract: We present a basic introduction to genetic terminology that has implications for growers and users of native 

plants. An understanding of underlying genetic concepts is essential for ensuring that plant material of an appropriate source 

is planted on restoration sites.

Link: http://npj.uwpress.org/content/5/2/105.full.pdf+html

609. Smith SL, Sher AA, Grant TAI. 2007. Genetic diversity in restoration materials and the impacts of seed collection 

in Colorado’s restoration plant production industry. Restoration 

Ecology 15:369-374.

Type: Journal 

Geographic Area: Colorado, USA

Keywords: ecotype, genetic diversity, native plant production, seed collection

Abstract: The ever increasing demand for native plants and 

seed for use in restoration and revegetation has created a sizable industry. The large scale production and planting of native 

plants have given rise to a suite of ecological concerns including collection impacts, genetic diversity, and provenance. This 

study examines the practices and beliefs of 12 restoration plant 

production companies in Colorado with regard to arising ecological issues and identifies where further research is needed. 

We found that native seed collection in Colorado was largely 

unregulated and unmonitored and impacts were unknown. 

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seed orchards. We use a literature review to evaluate current 

legislation and to develop recommendations for herbaceous 

and woody species. 

613. Willan RL, Barner H. 1993. Matching seed source 

to planting site. Humlebaek, Denmark: Danida Forest Seed 

Centre. Lecture Note B3. 25 p.

Type: Government Document

Geographic Area: Denmark, Europe

Compilers’ Keywords: restoration, seed transfer, local adaptation, seed provenance

Compilers’ Summary: This is a lecture on matching seed 

source to planting site from the Danida Forest Seed Centre 

in Denmark. It defines and discusses planting sites and seed 

sources in reference to considerations in reforestation efforts. 

Organized into six sections, it covers needs and methods for 

describing seed sources and planting sites, categories of sites 

and seed sources, problems in seed transfer of local and nonlocal sources, survey requirements, and use of improved seed 

sources (phenotypically or genetically) to new planting sites. 

Link: http://curis.ku.dk/ws/files/20656861/b3_002.pdf

614. Williams G. 2007. Local provenance plant seed and 

restoration: scientific imperative or romantic diversion? New 

South Wales, Australia. 20 p.

Type: Report

Geographic Area: Australia

Compilers’ Keywords: seed source, restoration

Introduction: It is a widely held assumption that plants 

used in revegetation projects should be derived from “local 

provenance” seed. Throughout the last twenty years or so 

this notion has been a central tenet of the extension literature 

in government funded natural resource management programs such as Landcare. On the whole I believe this assumption is based on unjustified and tenuous interpretations on 

the related scientific literature. Indeed in the last five years 

or so some individuals and organizations have begun to acknowledge that poor or even over-excited interpretation of 

provenance research has stymied good operational practice 

for ecological restoration. It is hoped that we are now seeing the beginning of the end of a simplistic application of 

the phenomenon of provenance to the practice of restoration. 

Nonetheless an insistence on using “local” seed retains a 

dominant place in the thinking of many practitioners within 

the natural resource management sector, contributing, in my 

view, to wasted resources and poor conservation outcomes. 

In this paper I argue that attempts at defining provenance in 

terms of fixed geography are unnecessary, technically unfeasible, and ultimately non-scientific, and that habitat matching 

is a better approach to identifying suitable sources of seed. I 

also attempt to use the discourse surrounding the promotion 

say in the following—studies have been made of a number of 

plant species. Usually species with extensive distribution, each 

occurring in as many different localities as possible, have been 

selected. The conditions in the natural habitats have been noted 

as far as possible, and seeds and pieces of turf have been collected and brought home from various habitats in order that 

the types might be followed in permanent cultures laid out in 

the experimental fields. A review at this stage of some of the 

results of these studies is intended to show the necessity and 

importance of a comparative study of organisms in their natural habitats and in the experimental field when the question of 

the differentiation of species and varieties is to be discussed.

612. Vander Mijnsbrugge K, Bischoff A, Smith B. 

2010. A question of origin: where and how to collect seed for 

ecological restoration. Basic and Applied Ecology 11:300-311.

Type: Journal 

Geographic Area: Global

Keywords: seed provenance, commercial seed collection, 

home-site advantage, local adaptation, habitat differentiation, 

seed collection zones, inbreeding and outbreeding depression, 

seed transfer, seed orchard 

Abstract: Native plant species are routinely planted or sown 

in ecological restoration projects, but successful establishment 

and survival depend on where and how seeds are collected. 

Research suggests that it is important to use locally adapted 

seeds. Local populations often show a home-site advantage 

and non-local genotypes may be maladapted to local environmental conditions. Furthermore, intraspecific hybridisation of 

local and non-local genotypes may have a negative impact on 

the genetic structure of local populations via mechanisms such 

as outbreeding depression. Many species show a strong smallscale genetic differentiation between different habitats so that 

matching habitats of the restoration and donor site can be more 

important than minimizing geographical separation. It is a 

challenge to identify appropriate seed sources because strong 

small-scale population differentiation makes it difficult to delineate geographically defined seed zones to which seed exchange should be limited. Moreover, it is important to consider 

the genetic diversity of introduced material because it may be 

crucial to avoid genetic bottlenecks, inbreeding depression and 

poor establishment of plant populations. Repeated propagation 

in stock, which is often required to obtain a sufficient amount 

of seeds, can further reduce genetic diversity and may select 

for particular genotypes. Negative impacts of improper seed 

choice for nursery planting stock may become detectable only 

after many years, especially in long-lived and slow growing 

plants. Although scientific information on many species remains limited, the increasing demand for translocation of seed 

means that mandatory regulations are necessary. Guidelines 

should prescribe a specification of seed provenance, a record 

of genetic diversity of wild collections and rules for subsequent 

processing such as direct transfer and propagation of stock or 

CHAPTER 4 • TRANSFER GUIDELINES AND ZONES • GENERAL
218 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

“native” and “local” have vague or imprecise meanings and 

are discussed in the context of plant selection.

Link: http://extension.oregonstate.edu/catalog/pdf/em/

em8885-e.pdf

616. Ying CC, Yanchuk AD. 2006. The development 

of British Columbia’s tree seed transfer guidelines: purpose, concept, methodology, and implementation. Forest Ecology and 

Management 227:1-13.

Type: Journal 

Geographic Area: Western Canada

Keywords: seed transfer, genecology, seed zones, climate 

change

Abstract: The development of forest tree seed transfer research, guidelines, regulations and policy has a long history in 

Canada, as well as in many other parts of the world. While the 

implicit assumptions of what is involved in developing seed 

transfer limits, guidelines and policy are generally accepted, 

the scientific and biological processes that underpin their validity are not readily available to most foresters. We provide an 

overview of the historical and current technical approaches to 

the development of seed transfer in British Columbia, and the 

overall framework which incorporates key biological, statistical and administrative issues in regulating the movement of 

forest tree seed. An example of how seed transfer information 

is developed from field experiments to guidelines or limits 

is provided from the lodgepole pine provenance tests in BC. 

Seed transfer research as it relates to the movement of wild 

or seed orchard seed will need to factor in the complications 

being predicted with climate change. As such, seed transfer 

research will continue to evolve as field experiments mature, 

new tests are established, statistical approaches and geographic 

information systems improve, and climate prediction tools attain greater resolution.

Link: http://www.abtreegene.com/toolkit/pdf/Adapt_Ying.

pdf

of provenance and local seed to address the broader idea of 

the relationship between culture, ecological knowledge and 

landscape-scale conservation “design.” The practice of ecological restoration represents a cultural aspiration that inevitably results in the creation of new kinds of ecosystems no 

matter how ecologically stringent or scientifically objective 

our criteria and strategies appear to be. The provenance issue 

sheds light on the negative impact that poorly thought out 

ideas and approaches in applied ecology have to progress 

in achieving restoration goals. This is particularly the case 

where there is a failure to candidly acknowledge our role 

as active landscape designers and managers, and agents of 

landscape change. 

Link: http://www.florabank.org.au/files/documents/provenance/Local_Provenance_Seed_Design_Romance_Science.

pdf

615. Withrow-Robinson B, Johnson R. 2006.

Selecting native plant material for restoration projects: ensuring 

local adaptation and maintaining genetic diversity. Corvallis 

(OR): Oregon State University Extension Service. EM 8885-

E. 10 p.

Type: Report

Geographic Area: Pacific Northwest USA

Compilers’ Keywords: common garden, ecoregion, seed 

zones, transfer guidelines

Introduction: Selecting appropriate plant materials for 

restoration projects contributes to the success of the project by ensuring better adaptation and survival of the plants. 

“Appropriate” means choosing species that are suitable for 

the site, are grown from locally adapted sources, and have a 

solid genetic base. This publication is for people involved in 

the important task of rehabilitating and restoring Oregon’s 

threatened or degraded habitats. Growing numbers of private 

landowners, local organizations, and government entities are 

embarking on projects to restore the health or function of 

their riparian areas, wetlands, prairies, savannas, and other 

habitats. Choosing the right plants is critical to the long-term 

success of these projects, but it is not always as straightforward as it seems. Managers often lack a strong background 

in genetics and may be confused by competing (and sometimes conflicting) claims when collecting or buying seeds, 

seedlings, or other plant materials. The purpose of this publication is to help restoration managers ask and respond to 

two important questions: 1. Where should our plants come 

from; or, more specifically, what are the appropriate sources 

of origin for the plants to be used in our project? 2. Are we 

maintaining adequate genetic diversity in the plant materials we introduce? In this publication, we describe important 

concepts, such as source of origin, and management mechanisms, such as transfer guidelines. (Terms in bold are in 

the glossary, page 9.) Other commonly used terms such as 

CHAPTER 4 • TRANSFER GUIDELINES AND ZONES • GENERAL
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 219

617. Adams WT, Zuo J, Shimizu JY, Tappeiner JC. 

1998. Impact of alternative regeneration methods on genetic diversity in coastal Douglas-fir. Forest Science 44:390-396.

Type: Journal 

Geographic Area: Oregon, USA

Keywords: Pseudotsuga menziesii var. menziesii (Mirb.) 

Franco, allozymes, artificial reforestation, natural regeneration, gene conservation

Abstract: Genetic implications of natural and artificial regeneration following three regeneration methods (group selection, shelterwood, and clearcut) were investigated in coastal 

Douglas-fir (Pseudotsuga menziesii var. menziesii [Mirb.] 

Franco) using genetic markers (17 allozyme loci). In general, 

harvesting followed by either natural or artificial regeneration 

resulted in offspring populations little altered from those in the 

previous generation. Cutting the smallest trees to form shelterwoods, however, resulted in the removal of rare, presumably deleterious, alleles, such that slightly fewer alleles per 

locus were observed among residual trees (2.76) and natural 

regeneration (2.75) than found in uncut (control) stands (2.86). 

Thus, although the shelterwood regime appears quite compatible with gene conservation, it would be best to leave parent 

trees of a range of sizes in shelterwoods designated as gene 

conservation reserves, in order to maximize the number of 

alleles (regardless of current adaptive value) in naturally regenerated offspring. Seedling stocks used for artificial regeneration in clearcut, shelterwood, and group selection stands (7 

total) had significantly greater levels of genetic diversity, on 

average, than found in natural regeneration. This is probably 

because the seeds used in artificial seedling stocks came from 

many wild stands and thus, sampled more diversity than found 

in single populations.

Link: http://www.fsl.orst.edu/pnwtirc/Electronic%20version%20of%20some%20pubs/Adams,%20Zuo,%20Shimizu,%20et%20al%20(1998).pdf

618. Andrew RL, Wallis IR, Harwood CE, Foley WJ. 

2010. Genetic and environmental contributions to variation 

and population divergence in a broad-spectrum foliar defense of 

Eucalyptus tricarpa. Annals of Botany 105:707-717.

Type: Journal

Geographic Area: Australia

Keywords: additive genetic variance, open-pollinated common-garden experiment, genotype × environment interaction, 

plasticity, Eucalyptus tricarpa, sideroxylonal, formylated phloroglucinol compounds (FPCs), chemical defence

General (583–616)—reviews, history, 

summaries, justification

Research (617–802)—genetic studies, tree 

improvement, climate change

Strategies (803–827)—framework, delineation, modelling

Resources (828–877)—guidelines, zones, 

maps, tools, software

Transfer Guidelines

and Zones

CHAPTER 4 • TRANSFER GUIDELINES AND ZONES • RESEARCH

#. Author(s), Date. Title. Source.

Type: 

Geographic Area: 

Keywords: 

Description:

Link:

location of study

keywords written by the 

author(s) or provided by the 

compilers

ID number and

information

abstract, summary, or 

portions of the introduction 

or conclusion written by the 

author(s) or provided by the 

compilers

book, book section, conference 

paper, dissertation, government 

document, journal article, 

newsletter, presentation, report, 

thesis or web page

internet address of reference

Each reference contains the following:
220 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

plant propagules used in revegetation projects are from a location that is geographically or ecologically similar to the planting site. Local adaptation and population genetic differentiation 

studies have provided evidence supporting the use of local plant 

materials for revegetation with several native plant species, primarily grasses. However, along with grass species, shrub species 

are also frequently used in revegetation seed mixes. To better 

understand the consequences of using non-local plant materials, I chose to study the population biology of the widespread 

shrub winterfat (Krascheninnikovia lanata, Chenopodiaceae) 

among five populations in New Mexico, USA. The investigation of winterfat population biology included a comparison of 

winterfat plant morphology In situ, quantification of the ecological distance between the five sites, and measurement of the rate 

of emergence and floral onset in a greenhouse common garden. 

I also carried out a reciprocal transplant experiment in which individuals from different locations were planted in replicate common gardens. Transplant survival, size, and reproduction were 

quantified for two years. To complement the morphological studies, the genetic structure of winterfat was quantified using nine 

isozyme loci. Genetic variation, population differentiation, and 

correlations among genetic, geographic, and ecological distance 

were assessed. The following questions were addressed: (1) Do 

winterfat populations differ in the vegetative or inflorescence 

size of individuals in the field? (2) Do winterfat populations 

differ in emergence phenology in the greenhouse? (3) Do winterfat populations differ in floral phenology in the greenhouse? 

(4) How do study sites differ in soil characteristics, climate, and 

plant community composition? (5) Do winterfat plants perform 

better than plants from other locations at their site of origin? (6) 

Was the weather at the planting locations during the experiment 

different from the historical climate at those locations? (7) How 

much neutral genetic variation do these populations of winterfat possess? (8) How is this neutral genetic variation partitioned 

among populations? (9) Is genetic distance correlated with geographic or ecological distance? The data from these experiments 

will assist restoration practitioners in determining appropriate 

plant material sources for revegetation project.

620. Barnett JP, Brissette JC. 2007. Regenerating 

shortleaf pine: results of a 5-year cooperative research initiative. 

In: Kabrick JM, Dey DC, Gwaze D, editors. Shortleaf Pine 

Restoration and Ecology in the Ozarks. Newtown Square (PA): 

USDA Forest Service, Northern Research Station. General 

Technical Report NRS-P-15. p 105-111.

Type: Government Document

Geographic Area: Southern USA

Compilers’ Keywords: shortleaf pine, Pinus echinata, reforestation, transfer guidelines

Abstract: Shortleaf pine (Pinus echinata Mill.) is unique 

among the southern pines. It has the widest natural 

range and thrives on shallow rocky soils of the Interior 

Highlands, where most other pine species perform poorly. 

Abstract: Both environmental and genetic effects contribute 

to phenotypic variation within and among populations. Genetic 

differentiation of quantitative traits among populations has been 

shown in many species, yet it can also be accompanied by other 

genetic changes, such as divergence in phenotypic plasticity and 

in genetic variance. Sideroxylonal (a formylated phloroglucinol 

compound or FPC) is an important chemical defense in eucalypts. The effect of environmental variation on its production is 

a critical gap in our understanding of its genetics and evolution. 

The stability of genetic variation in sideroxylonal was assessed 

within and among populations of Eucalyptus tricarpa in three 

replicated provenance/progeny trials. The covariance structure 

of the data was also modelled to test whether genetic variances 

were consistent among populations and Fain’s test was applied 

for major gene effects. A significant genotype x environment interaction occurred at the level of population, and was related to 

temperature range and seasonality in source populations. Withinpopulation genetic variation was not affected by genotype × environment effects or different sampling years. However, withinpopulation genetic variance for sideroxylonal concentration 

differed significantly among source populations. Regression 

of family variance on family mean suggested that this trait is 

subject to major gene effects, which could explain the observed 

differences in genetic variances among populations. These results highlight the importance of replicated common-garden 

experiments for understanding the genetic basis of population 

differences. Genotype × environment interactions are unlikely to 

impede evolution or responses to artificial selection on sideroxylonal, but the lack of genetic variation in some populations may 

be a constraint. The results are broadly consistent with localized 

selection on foliar defence and illustrate that differentiation in 

population means, whether due to selection or to drift, can be accompanied by changes in other characteristics, such as plasticity 

and genetic variance.

Link: http://www.ncbi.nlm.nih.gov/pubmed/20228089

619. Barnes MG. 2009. The effect of plant source 

location on restoration success: a reciprocal transplant experiment with winterfat (Krasheninnikovia lanata) [Dissertation].

Albuquerque (NM): University of New Mexico. 156 p.

Type: Dissertation

Geographic Area: New Mexico, USA 

Compilers’ Keywords: common garden, genetic differentiation, In situ, morphological variation

Abstract: Ecological restoration is becoming more frequent 

due to the increased pace of land disturbance, more comprehensive government regulations, and the recognition of the valuable ecosystem services that natural areas provide. One part of 

restoration is revegetation, or the introduction of off-site plant 

materials to the restoration site. As the applications of revegetation have become more diverse, so too have the objectives of 

these projects. More specifically, it is increasingly important that 

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 221

growing seasons, and to moderately or extremely low winter temperatures. 

622. Benowicz A, Hirondelle SL, El-Kassaby YA. 

2001. Patterns of genetic variation in mountain hemlock (Tsuga 

mertensiana (Bong.) Carr.) with respect to height growth and frost 

hardiness. Forest Ecology and Management 154:23-33.

Type: Journal 

Geographic Area: Western Canada

Keywords: Tsuga mertensiana, intra-specific variation, frost 

hardiness, growth rate, seed transfer

Abstract: Genetic structure (variance among and within populations) and geographic pattern of variations in growth and frost 

hardiness of mountain hemlock (Tsuga mertensiana (Bong.) 

Carr.) populations from coastal British Columbia (BC) were examined. Populations fell into two main groups—south coast (16 

sources) and north coast (two sources). Height growth of greenhouse grown seedlings was measured during the first growing 

season on a weekly basis whereas frost hardiness and branch 

water content were evaluated during the fall at monthly intervals. Significant differences among the populations were found 

in maximum instantaneous growth rate, height at the end of the 

growing season and in October and November front hardiness. 

There were no differences in date of maximum growth rate, 

branch water content and September frost hardiness. Most of the 

genetic variance was found within populations: 77% for height, 

maximum growth rate and November frost hardiness and 87% 

for October frost hardiness. Geographic trends were identified 

using multiple linear regression and canonical correlation analyses based on population means and latitude, longitude and elevation of the population origin. The uneven distribution of sources 

limited the extrapolation, but some general trends appeared. 

Maximum instantaneous growth rates increased with latitude, 

elevation and decreased longitude (R2 = 0.64) and consequently 

were negatively correlated with the length of the growing season in the seed collection sites. Both populations from the north 

coast developed frost hardiness earlier than any population from 

the south coast. The relationship between climate coldness and 

growth rates has potential implications for seed transfer guidelines. Due to their higher growth rates, plants from higher altitudes may accomplish the same amount of growth within a 

shorter period compared to plants from lower altitudes when 

both are transferred north. Results of the frost hardiness tests 

indicate that seed transfer along the British Columbia coast 

of more than 3 degrees northward will considerably increase 

chances of frost damage in plantations. 

Link: http://www.researchgate.net/publication/222958778_Patterns_of_genetic_variation_in_mountain_hemlock_(Tsuga_mertensiana_(Bong.)_Carr.)_

with_respect_to_height_growth_and_frost_hardiness/

file/79e41513134e126d13.pdf

Although wood quality is excellent, it has been one of the 

most neglected species from both research and operational 

standpoints. It has a history of poor performance following 

outplanting with survival of less than 50%. The technology to change this situation was developed after formation 

of the Shortleaf Pine Artificial Regeneration Taskforce in 

1984. Over a 6-year period, 15 studies were installed in 

Arkansas and Oklahoma to address seedling production 

and establishment. Information resulting from these studies resulted in increased seedling survival in both the Ozark 

and Ouachita National Forests. This paper summarizes research from these and other studies that led to the improved 

success in reforestation of the species.

Link: http://www.treesearch.fs.fed.us/pubs/12840

621. Bates CG. 1930. The frost hardiness of geographic 

strains of Norway pine. Journal of Forestry 28:327-333.

Type: Journal 

Geographic Area: Central USA

Compilers’ Keywords: geographic strains

Introduction: Norway pine (Pinus resinosa) is a species 

which covers a comparatively narrow latitudinal range, although in its range from the northeastern coast to the Lake 

States and southern Canada it encounters summer temperature differences of about 10 °F and considerably greater differences in mid-winter temperatures (-35 and -40 °F in the 

northwestern part of the range to not much below zero in 

the Alleghenies, these being mean annual minima). Nearly 

as great differences are found in winter if mean January 

temperatures be considered, of from 0 to 30 °F. Because of 

its great commercial value and its extensive use on reforestation projects, Norway pine has been chosen by the Lake 

States Forest Experiment Station as the first species to be 

subjected to a scrutinizing study of geographical, varietal, 

and individual differences, or in other words to a “breeding” study whose primary purpose is to determine what 

“seed zones” should be recognized in order to avert failures in planting due to lack of local adaptation. But because 

of the great uniformity of appearance and development of 

the species, as well as the considerations mentioned in the 

first paragraph above, the writer has felt some doubts as to 

whether outstanding differences would be likely to be developed by such a study. Therefore, to “anticipate” to some 

extent the results of field comparisons which were started 

at the same time through nursery sowings of 41 different 

collections of Norway pine seed, an indoor experiment was 

begun which it was hoped would bring out the existence of 

physiological differences affecting hardiness. Without going into the question of what comprises hardiness to freezing temperatures, it may be stated as more or less obvious 

that differences within a species should develop according 

as its local forms have become adapted to long or short 

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222 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

for the 2 hybrids. It was demonstrated by studying correlation 

between parent phenotype and parent crossbreeding value in 

E. urophylla that the higher additive variance for females was 

probably due to selection of males for their phenotypic value. 

This result implies a weaker selection intensity for the males in 

order not to heavily reduce the variability. Variance structure 

was different according to considered hybrids. For circumference, additive variance accounted for 80% of total genetic variance in E. urophylla • grandis and only 40% in E. urophylla • 

pellita. Sampling of parents and/or genetic distance between 

species could be an explanatory factor of this difference.

Link: http://www.silvaegenetica.com/fileadmin/content/dokument/archiv/silvaegenetica/45_1996/45-2-3-171.pdf

625. Bower AD, Aitken SN. 2006. Geographic and 

seasonal variation in cold hardiness of whitebark pine.

Canadian Journal of Forest Research 36:1842-1850.

Type: Journal 

Geographic Area: California, USA

Compilers’ Keywords: Pinus albicaulis, common garden, 

adaptation

Abstract: Artificial freeze-testing utilizing the electrolyteleakage method was used to test the cold hardiness of 2-yearold whitebark pine (Pinus albicaulis Engelm.) seedlings growing in a common garden. Testing across all seasons was used to 

determine the annual pattern of cold hardiness, and more intensive sampling in the fall and spring was used to assess genetic 

variation in cold injury among geographic regions spanning the 

range of the species. Mean hardiness varied widely from -9 

°C in early summer to below -70 °C in the winter. Trees from 

interior and northern regions were the hardiest in the fall, while 

trees from California were the least hardy. Geographic patterns 

of hardiness in the spring were reversed. Significant differences 

in cold injury among regions were detected on all dates except 

during the winter. Heritability was low to moderate for both 

the spring (h2 = 0.18) and the fall (h2 = 0.28), and genetic correlation was weak (rA = 0.18). Only spring cold injury was genetically correlated with date of needle flush (rA = 0.34). Mean 

cold injury in the fall was most closely correlated with mean 

temperature of the coldest month in the parental environment (r 

= 0.81). Whitebark pine is well adapted to the low temperatures 

of the harsh environments where it is found; however, regional 

variation indicates that moving seed for restoration purposes 

from areas with higher winter temperatures to colder environments may increase the chance of fall cold injury.

626. Bower AD, Aitken SN. 2007. Genetic diversity and 

geographic differentiation in quantitative traits and seed transfer guidelines for whitebark pine. In: Goheen EM, Sniezko R, 

editors. Whitebark Pine: A Pacific Coast Perspective. Portland 

(OR): USDA Forest Service, Pacific Northwest Region. R6-NRFHP-2007-01. p 98-101.

623. Bhattarai K, Bushman BS, Johnson DA, 

Carman JG. 2010. Phenotypic and genetic characterization of western prairie clover collections from the western United 

States. Rangeland Ecology & Management 63:696-706.

Type: Journal 

Geographic Area: Western USA

Keywords: amplified fragment length polymorphism (AFLP), 

conservation, Dalea, Great Basin, legume, population structure, 

revegetation

Abstract: Few North American legumes are available for 

rangeland revegetation in the semiarid western United States. 

Western prairie clover (Dalea ornata [Douglas ex Hook.] Eaton 

& J. Wright) is a perennial legume with desirable forage characteristics and is distributed in the northern Great Basin, Snake 

River Basin, and southern Columbia Plateau. Understanding 

the genetic and ecotypic variability of this species is a prerequisite for developing populations suitable for revegetation 

purposes. To address this need, we established two commongarden plots of western prairie clover from 22 sites in Idaho, 

Oregon, and Washington. Significant variation was detected 

among the collections for all traits measured. Among the measured traits, flowering date was correlated with collection-site 

temperature and elevation. Population structure estimates from 

474 amplified fragment length polymorphism markers resulted 

in two distinct, genetically differentiated groups and a third 

admixed group, and flowering date played a significant role in 

discriminating those genetic-based groupings of collections. 

Positive correlations were observed between phenotypic and 

genetic distance matrices (r = 0.33, P = 0.005), phenotypic and 

geographic distance matrices (r = 0.35, P = 0.002), and genetic 

and geographic distance matrices (r = 0.31, P = 0.009). Based 

on these results, we recommend that two germplasm sources of 

western prairie clover be developed for use across the collection 

area, one from the Deschutes River region and the other encompassing Idaho, Washington, and eastern Oregon collection sites.

624. Bouvet JM, Vigneron P. 1996. Variance structure 

in Eucalyptus hybrid populations. Silvae Genetica 45:2-3.

Type: Journal 

Geographic Area: Congo, Africa

Compilers’ Keywords: reciprocal recurrent selection, factorial mating design, variance components, Eucalyptus urophylla, 

Eucalyptus pellita, Eucalyptus grandis

Summary: This article presents the analyses of the first mating 

designs established in the reciprocal recurrent selection scheme 

of Eucalyptus in Congo. Two selection schemes were involved, 

the first one concerning E. urophylla • grandis, the second one 

concerning E. urophylla • pellita. For each of the hybrid species, 

height, circumference and volume were measured until 3 of 4 

years (half of the rotation age). Male additive variance was lower than female additive variance (30% of the additive variance) 

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 223

from 48 populations were grown for two years and measured 

for height increment, biomass, root to shoot ratio, date of needle flush, fall and spring cold injury, and survival. Significant 

variation was observed among populations for most traits. 

The QST was low (0.07–0.14) for growth traits and moderate 

(0.36–0.47) for cold adaptation related traits, but varied by region. Cold adaptation traits were strongly correlated with mean 

temperature of the coldest month of population origins, while 

growth traits were generally correlated with growing season 

length. We recommend that seed transfer for restoration favor 

seed movement from milder to colder climates to a maximum 

of 1.9 °C in mean annual temperature in the northern portion 

of the species range, and 1.0 °C in the U.S. Rocky Mountains 

to avoid maladaptation to current conditions yet facilitate adaptation to future climates.

628. Broadhurst LM, North T, Young AG. 2006. 

Should we be more critical of remnant seed sources being used 

for revegetation? Ecological Management and Restoration 

7:211-217.

Type: Journal 

Geographic Area: Australia

Keywords: Acacia, fitness, fragmentation, genetics, restoration, 

revegetation, seed quality, self-incompatibility

Abstract: A challenge for land managers restoring degraded 

agricultural landscapes across southern Australia will be to 

ensure the viability of remnant vegetation while simultaneously supplying the quantities of appropriate seed required for 

revegetation. To ensure such revegetation programs have the 

best chance of success, seed that is both genetically diverse 

and locally adapted will be required. Identifying suitable seed 

sources can be particularly difficult in regions where local seed 

sources are restricted to small and isolated remnants. Golddust Wattle (Acacia acinacea Lindl.) is a key revegetation species in the Deniliquin region of New South Wales; however, 

broadscale land clearing in the area has limited local seed 

sources to a few remnant stands. Field-based experience suggests that revegetation success may depend upon the source of 

seed used, raising the question of whether differences in the 

germination and survival of seed reflect functional problems 

within these source populations. To test this possibility, seed 

was collected from 15 mothers in each of three seed sources 

regularly used for local restoration programs. Seed quality 

for each mother was assessed in terms of seed production 

and seedling fitness. In addition, genetic diversity and mating 

system parameters were determined to assess whether these 

explained the seed quality responses observed. Differences 

among the seed sources with respect to seed quality were generally congruent with field-based predictions. High levels of 

correlated paternity in the two poorly performing seed sources 

probably reflect limited mating availability due to smaller 

population sizes and genetic incompatibility being mediated 

by a self-incompatible reproductive strategy. Further research 

Type: Government Document

Geographic Area: Western Canada

Compilers’ Keywords: Pinus albicaulis, restoration, common 

garden, white pine blister rust, mountain pine beetle, Dedroctonus 

ponderosae

Introduction: Whitebark pine (Pinus albicaulis Englem.) has 

declined dramatically throughout its range due to white pine 

blister rust (caused by the fungus Cronartium ribicola J.C. 

Fisch.), successional replacement resulting from fire suppression, and attack by mountain pine beetle (Dendroctonus ponderosae Hopkins). Restoration is needed to halt or reverse this 

decline; however, no information regarding genetic diversity 

and local adaptation of quantitative traits is available to guide 

these efforts. A seedling common garden experiment was employed to assess genetic diversity and geographic differentiation 

of quantitative traits (QST) of whitebark pine and to determine 

the climatic variables driving local adaptation. Seedlings from 

48 provenances from a near range-wide seed collection were 

grown in raised beds in Vancouver, B.C. for two years in two 

soil temperature treatments (ambient and cold). Seedlings were 

measured for second year height increment, total biomass, root : 

shoot ratio, date of needle flush, fall and spring cold injury, and 

survival. Significant differences were found between soil temperature treatments for height growth and survival, with seedlings in the cold treatment performing better. The environment 

where the test was grown (Vancouver, B.C.) is considerably 

warmer than the natural habitat of whitebark pine. While most 

temperate forest trees would likely benefit from the warmer 

soil temperatures, it appears that this is a stressor for a species 

adapted to cold, harsh environments.

Link: http://www.fs.fed.us/outernet/r6/nr/fid/wbpine/

papers/2007-wbp-wpbr-resist-bower.pdf

627. Bower AD, Aitken SN. 2008. Ecological genetics and seed transfer guidelines for Pinus albicaulis (Pinaceae). 

American Journal of Botany 95:66-76.

Type: Journal 

Geographic Area: Western North America

Keywords: genetic variation, geographic differentiation, local 

adaptation, Pinus albicaulis, quantitative traits, seed transfer, 

whitebark pine, white pine blister rust

Abstract: Whitebark pine (Pinus albicaulis Engelm.) has 

greatly declined throughout its range as a result of introduced 

disease, fire suppression, and other factors, and climate change 

is predicted to accelerate this decline. Restoration is needed; 

however, no information regarding the degree of local adaptation is available to guide these efforts. A seedling commongarden experiment was employed to assess genetic diversity 

and geographic differentiation (QST) of whitebark pine for 

traits involved in growth and adaptation to cold and to determine climatic variables revealing local adaptation. Seedlings 

CHAPTER 4 • TRANSFER GUIDELINES AND ZONES • RESEARCH
224 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Introduction: ‘Seed transfer’ in reforestation is the process 

of moving seed, or seedlings, from place of origin to alternative planting sites. Strong support for the theory of optimality 

of local populations has been presented by Langlet (1936) and 

Clausen, Keck & Hiesey (1948). Consequently, use of nonlocal seed is avoided if possible to minimize losses of vigor or 

fitness; but in cases where local seed is not available, transfer 

guides are necessary. For Douglas-fir (Pseudotsuga menziesii

(Mirb.) Franco) in western Washington and Oregon, present guides are rules of thumb taken from Swedish models. 

Therefore, a method for early examination of effects of moving 

seed would be helpful. This report presents a simple predictive 

model which uses timing of vegetative bud burst for examining responses of seedlings to transfer. The main assumptions 

included in the model were: (1) genetic differentiation in timing of bud burst is in response to adaptive selection; (2) the 

timing of bud burst in an adapted phenotype is a function of 

the physiological response of that provenance to temperature; 

(3) chilling requirements of all provenances will be satisfied 

by normal winter temperatures, regardless of the part of the 

region to which they are moved. Only the first assumption is 

examined in this report, by the method of provenance- habitat correlations. The other two assumptions are supported in 

the literature; for example, timing of bud burst appears to be 

almost exclusively temperature mediated if chilling requirements are met (Wommack 1964; Lavender & Hermann 1970). 

Also the chilling requirement of Douglas-fir is less than 80 

days of temperatures lower than 10 °C (Wommack 1964), 

whilst in any winter most of the region under consideration 

will have more than 100 days in which maximum temperatures 

will not rise above 120 °C (Wakefield 1969). The exceptions 

are in south-coastal Oregon and California, a negligible part of 

the Douglas-fir region. The model required two sets of equations. The first, based on an experiment using forty-four provenances, describes timing of bud burst as a function of average 

daily temperature. The second describes pattern of average 

daily temperature at stations where provenances were collected. Together, the equations provided functions for predicting bud-burst dates. Predicted dates were then used to examine 

hypothetical effects of transferring seed along elevational and 

latitudinal gradients.

631. Campbell RK. 1979. Genecology of Douglas-fir in a 

watershed in the Oregon Cascades. Ecology 60:1036-1050.

Type: Journal 

Geographic Area: Oregon, USA

Keywords: adaptation, Douglas-fir, genetic differentiation, genetic variance, natural selection, provenance, seed source, seed 

transfer, topocline

Abstract: To gain insight into genetic microstructure of subregional populations of coastal Douglas-fir (Pseudotsuga menziesii

[Mirb.] Franco var. menziesii), genetic variability in a population found on a 6,100-ha, central Oregon watershed is described. 

is now required to determine whether observed variability 

in the quality of seed from remnant vegetation in degraded 

landscapes is compromising revegetation efforts, and to help 

practitioners develop strategies to critically evaluate their seed 

sources.

629. Broadhurst LM, Young AG, Thrall PH, Murray 

BG. 2006. Sourcing seed for Acacia acinacea, a key revegetation 

species in South Eastern Australia. Conservation Genetics 

7:49-63.

Type: Journal 

Geographic Area: Australia

Keywords: AFLP, morphology, ploidy, revegetation, seed 

sourcing

Abstract: Intensive large-scale revegetation programs are 

a major activity in degraded landscapes. The collection 

and deployment of seed for revegetation is primarily based 

on local seed sources to preclude concerns associated with 

local adaptation as well as outbreeding depression in small 

remnant populations. For most species however, little is 

known about the levels of genetic diversity being sourced 

and deployed, or even of the levels of adaptively significant variation present across the species range. Acacia 

acinacea (Gold Dust wattle) is a key revegetation species 

widely distributed across south eastern Australia. Levels 

of variation in terms of morphology, fitness, ploidy level 

and AFLPs were assessed using seed lots from 35 populations across the Murray Darling Basin in south eastern 

Australia. Multivariate analysis indicated that three of the 

populations were misidentified while the remaining populations were differentiated into three groups. One of these 

groups (Bendigo) was highly geographically localized and 

tetraploid while the other two (Group 1 and Group 2) were 

both diploid with overlapping distributions. Fitness differences were also evident among these three groups while 

AFLPs indicated that they were genetically differentiated 

and that lower levels of diversity were captured from some 

populations. A major finding of this study was that no one 

technique was able to fully describe the variation present 

in A. acinacea and seed sourcing guidelines based on any 

single approach could have resulted in erroneous decisions being made. Information generated from this study 

has been used to refine seed sourcing guidelines.

630. Campbell RK. 1974. Use of phenology for examining 

provenance transfers in reforestation of Douglas-fir. Journal of 

Applied Ecology 11:1069-1080.

Type: Journal 

Geographic Area: Pacific Northwest USA

Compilers’ Keywords: Pseudotsuga menziesii, seed transfer 

guideline, genetic differentiation, adaptive phenotype

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the conclusion is that for genetically heterogeneous species in 

mountainous regions, genetic mapping is a prerequisite to directly estimating transfer effects by long-term tests. 

Link: http://www.silvaegenetica.com/fileadmin/content/dokument/archiv/silvaegenetica/35_1986/35-2-3-85.pdf

633. Campbell RK. 1991. Soils, seed-zone maps, and 

physiography: guidelines for seed transfer of Douglas-fir in 

southwestern Oregon. Forest Science 37:973-986.

Type: Journal 

Geographic Area: Oregon, USA

Keywords: Pseudotsuga menziesii, adaptation, ecogeographic 

center-genetic variation, genecology

Abstract: One procedure for guiding seed transfer is to partition the species habitat into zones within which there is little 

genetic variation from location to location. This report compares 

soil types and the existing regional seed zones as bases for classifying geographic genetic variation into zones. The data used 

were genotypic values of 135 Douglas-fir (Pseudotsuga menziesii) trees from 80 locations in southwestern Oregon. Genotypic 

values were estimated by measuring traits expressing phenology and growth potential of 2-yr seedlings in a common-garden 

experiment. Neither the soils model nor the seed-zones model 

satisfactorily classified geographic genetic variation (significant 

lack of fit). When physiographic variables (latitude, longitude, 

elevation, etc.) were added to the models, the added variables 

accounted for 16% to 23% of the total variation among locations. Gradients of genetic variation therefore exist within soil 

types and present seed zones. These gradients reflect the overall 

gradients of geographic variation with latitude, longitude, and 

elevation that occur in the region. The results suggest that zones 

constructed from physiographic model may explain more of the 

genetic variation in southwestern Oregon than do either soils or 

seed-zones models.

634. Campbell RK, Sorensen FC. 1978. Effect of test 

environment on expression of clines and on delimitation of seed 

zones in Douglas-fir. Theoretical Applications of Genetics 

51:233-246.

Type: Journal 

Geographic Area: Pacific Northwest USA

Keywords: Pseudotsuga menziesii, genecology, provenance, 

genotype-temperature interactions, fertilizer interactions

Abstract: Clinal models of population structure in an indigenous tree species can be used to delineate seed collection zones and breeding zones, and to devise transfer rules. 

Models may be developed by growing populations in test 

environments; however, a clinal description may be a function of test environments as well as of population genotypes. 

This possibility was studied by growing seedlings from 40 

populations of northwestern U.S. Douglas-fir (Pseudotsuga 

Genotypic values of 193 parent trees located throughout the 

watershed were estimated from progeny grown in a common 

garden. Then, genetic variation was partitioned into components 

attributable to parent-tree location and to differences among 

trees within locations. Within-location variation appeared to 

be homogeneous in the watershed; between-location variation 

was related to topography, but the patterns of trait variation differed depending on the trait measured. Growth traits exhibited 

strong gradients with elevation, but the relationship varied depending on position within the watershed. Based on estimates 

of proportions of nonadapted seedlings in hypothetical transfers 

of seed from one part of the watershed to another, genetic differentiation within the watershed was surprisingly large. In one 

transfer of 3.5 km between north- and south-facing slopes at the 

same elevation: 80% of seedlings were estimated to be poorly 

adapted. Although other possibilities exist, the topoclinal variation in traits probably results from selection as does the large 

within-location variation. Both kinds of variation are due to high 

selection intensities in the seedling stage, the former to selection by average environmental differences along gradients, the 

latter to microenvironmental heterogeneity. The combination of 

high within- and between-habitat variation is suited to a species 

which episodically colonizes an environment that is extremely 

heterogeneous in time and space.

Link: http://andrewsforest.oregonstate.edu/pubs/pdf/

pub1701.pdf

632. Campbell RK. 1986. Mapped genetic variation of 

Douglas-fir to guide seed transfer in southwest Oregon. Silvae 

Genetica 35:2-3.

Type: Journal 

Geographic Area: Oregon, USA

Compilers’ Keywords: seed source, seed transfer zones, provenance, Pseudotsuga menziesii

Summary: A procedure is illustrated for using mapped genetic 

variation in indigenous species to develop provisional seed 

transfer rules and seed zones. Genotypic values for 13 traits of 

135 parent trees from 80 locations furnished data for Douglasfir in a region in southwest Oregon. Values were estimated 

from open-pollinated progeny grown in two nursery beds. 

Two principal components accounted for about 96% of the 

total family and seed-source variation in all traits. There were 

strong gradients with location variables: elevation, latitude, 

distance from the ocean, slope, and sun exposure as affected by 

shade of adjacent mountains. Seed transfer rules and a procedure for calculating relative risk indicated that risks were large 

when seed was transferred either east-west along the southern 

boundary or north-south along the western boundary of the 

region. These gradients in risk coincide with the steepest precipitation and temperature gradients in the region. Advantages, 

disadvantages, and potential sources of error in the procedure 

are discussed. In spite of the limitations of genetic mapping, 

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styroblock at Kalamalka had the least uprooting resistance and 

root biomass of all treatments. In addition, the relationship 

between uprooting resistance and ground line diameter was 

different between nurseries. The results of this study suggest 

that regardless of height and diameter leaving the nursery, seed 

source root resistance and growth continued to be impacted by 

nursery culture 5 years after planting.

636. Campbell KA, Pawuk WA, Harris AS. 1989.

Microgeographic genetic variation of Sitka spruce in southeastern 

Alaska. Canadian Journal of Forest Research 19:1004-1013.

Type: Journal 

Geographic Area: Alaska, USA

Compilers’ Keywords: Sitka spruce, Picea sitchensis, common garden, principal component analysis, seed transfer, 

microgeographic

Abstract: Microgeographic genetic variation among populations of Sitka spruce on Mitkof Island in southeastern Alaska 

is described. In two common-garden environments, we evaluated genotypes of 208 parent trees from 114 locations in a 

17,000-ha area. Two principal components accounted for most 

of the variation among locations in 11 traits measured to evaluate growth vigor and rhythm of 2-yr old seedlings. Regression 

analyses of factor scores derived from principal components 

revealed genetic gradients associated with elevation, slope, aspect, and west-east and north-south directions. Large amounts 

of additive genetic variation in factor scores occurred among 

trees within locations. When this variation within locations 

was used as a scale, variation among locations was also large. 

In an extreme case, locations differed in factor scores of the 

first principal component by about 3.0 units of the standard 

deviation of additive genetic variation in factor scores. Of the 

total differentiation in this case, elevational range (600 m) contributed 0.7 units of standard deviation, aspect contributed 0.9 

units, and distance (16 km) from north central to southeastern 

parts of the island contributed 1.4 units.

637. Carles S, Lamhamedi MS, Beaulieu J, Stowe 

DC, Margolis HA. 2011. Differences in growth and mineral 

nutrition of seedlings produced from ten white spruce seed 

orchards. New Forests 42:195-214.

Type: Journal 

Geographic Area: Eastern Canada

Keywords: Picea glauca, seed orchard, seedlings, logistic function, mineral nutrition, seed zones

Abstract: To meet the needs for improved spruce seeds in the 

province of Quebec, Canada, 17 first-generation white spruce 

seed orchards (SO) were established. These SOs are located 

in different bioclimatic domains and contain seed trees originating from geographically and genetically distinct sources. 

To evaluate the influence of SO on seedling growth and 

menziesii var. ‘menziesii’ [Mirb.] Franco) in eight nursery-bed 

treatments which contrasted air and soil temperatures and nutrition. Growth traits measured were stem diameter, top height, 

and dry weight; phenological traits were bud-burst and budset dates, extension period, and extension period midpoint. 

Population samples interacted significantly with soil temperature for growth traits, and with soil and air temperatures combined for phenological traits. Interactions were at least partly 

explained by complex clinal associations of seedling performance with elevation, with latitude, and with distance from 

the ocean of the populations sampled. Both the complexity and 

the gradient of the clinal pattern depended on the trait and on 

the specific test environment. The clinal patterns of greatest 

complexity were expressed in warm air and soil treatments. 

Dry top-weights of population samples were associated with 

latitudes for samples grown in warm soils, but this relationship was not apparent in cool soils. A discrepancy in bud-burst 

dates between extreme coastal and more inland populations 

was greatest in warm soil-warm air treatments and was negligible in cool soil-cool air treatments. Populations x temperature interactions were attributed to the differential response of 

population samples to spring temperature and photoperiod. It 

is proposed that first attempts at devising a model can be based 

on nursery or growth chamber tests, and that test environments 

should stress contrasting photo- and temperature-regimes. The 

estimate of clinal structure in Douglas-fir suggests that there 

is more risk within northwestern U.S. in moving provenances 

east-west than north-south, that this risk increases with elevation of provenances, and that north-south transfers are more 

critical near the coast than inland.

635. Campbell KA, Hawkins CDB. 2004. Effect of 

seed source and nursery culture on paper birch (Betula 

papyrifera) uprooting resistance and field performance.

Forest Ecology and Management 196:425-433.

Type: Journal 

Geographic Area: Western Canada

Keywords: root strength, paper birch, nursery practices, stocktype, seed source, ground line diameter

Abstract: Little is known about the effect of seed source and 

nursery culture on the growth and performance of artificially 

regenerated paper birch planted in the field. In the summer 

of 2000, trees were vertically uprooted in Prince George, BC 

using a tripod and winch device to determine if seed source 

and nursery culture significantly affected the root strength 

of 5-year-old paper birch. Trees from a seed source in the 

Skeena region had the greatest uprooting resistance, and the 

most root biomass, but the least root strength compared to the 

other three seed sources tested. The Skeena trees may be exhibiting an adaptive strategy which provides maximum root 

reinforcement across a wide range of environments. Uprooting 

resistance was similar for the 415D stocktypes grown at Red 

Rock and Kalamalka nurseries. However, trees grown in 515A 

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639. Chen THH, Howe GT, Bradshaw HDJ. 2002.

Molecular genetic analysis of dormancy-related traits in 

poplars. Weed Science 50:232-240.

Type: Journal 

Geographic Area: USA

Keywords: black cottonwood, Populus trichocarpa Torr. & 

Gray, eastern cottonwood, Populus deltoids Bartr. Ex Marsh.

Abstract: We studied the molecular genetics of dormancyrelated traits in an F2 family of poplar (Populus) hybrids derived 

from a cross between a northern genotype of black cottonwood 

and a southern genotype of eastern cottonwood by mapping 

quantitative trait loci (QTLs) and candidate genes. Dormancyrelated traits included timing of vegetative bud set, fall frost 

damage, chilling response, timing of vegetative bud flush, and 

winter survival under field conditions, as well as photoperiodic 

responses (timing of bud set and number of new leaves) in a 

warm greenhouse under either a uniform 8-h photoperiod or 

a naturally shortening photoperiod in the fall. QTL analyses 

were conducted using a linkage map consisting of AFLP, 

microsatellite, and candidate gene markers. The candidate 

genes were chosen because of their potential roles in either 

photoperiodic perception (PHYB1, PHYB2) or abscisic acid 

signal transduction (ABI1B, ABI1D, ABI3). Significant QTLs 

were detected for all dormancy-related traits, except for winter 

survival, which had a relatively low heritability compared with 

the other traits. Interestingly, half of the field bud set QTLs did 

not map near photoperiodic QTLs. This is consistent with the 

moderate genetic correlation between these traits (0.53 to 0.60) 

and suggests that genetic differences in photoperiodic responses 

play only a modest role in explaining genetic differences in the 

timing of bud set under field conditions. Except for ABI1D, 

each of the candidate genes tested mapped near one or more 

of the dormancy-related QTLs. We conclude that molecular 

markers and QTL analyses can be used to study the genetics 

of dormancy-related traits, to design more effective breeding 

programs, and to provide new insights into tree physiology.

640. Conkle MT. 1973. Growth data for 29 years from the 

California elevational transect study of ponderosa pine. Forest 

Science 19:31-39.

Type: Journal 

Geographic Area: California, USA

Keywords: Pinus ponderosa Laws., elevational transect, genetic 

adaptation, genotype and environmental interaction

Abstract: Ponderosa pine progenies from parents restricted 

in latitude but spanning 7,000 feet of elevation show significant growth differences in plantations at low-, mid-, and 

high-elevation test sites. At low- and mid-elevation sites tree 

heights and diameters of progenies from high-elevation parents were the smallest; those from the low-elevation parents, 

intermediate; and those from mid-elevation seed parents, 

morphology, seedlings produced with seeds originating from 

the ten most commonly used first-generation white spruce SO 

in Quebec were raised under similar nursery conditions. Tissue 

nutrient concentrations of seedlings evolved similarly among 

seed orchards during the second growing season. At the end 

of the growing season, only shoot phosphorus concentrations 

were significantly different. When modeled with a logistic 

function, there was a significant difference between height and 

diameter growth curve parameters of seedlings from distinct 

SO during the second growing season. These differences led to 

significant differences in the height and shoot dry mass of the 

seedlings at the end of their second growing season, but not in 

their diameter or root dry mass. The ten SOs were clustered in 

two groups according to the above-ground characteristics of 

their progeny. This limited amount of morphological diversity 

suggests that expanding the size of the present seed zones may 

be an appropriate course of action for white spruce seed production in Quebec.

638. Casler MD, Stendal CA, Kapich L, Vogel KP. 

2007. Genetic diversity, plant adaptation regions, and gene pools 

for switchgrass. Crop Science 47:2261-2273.

Type: Journal 

Geographic Area: Central and Eastern USA

Compilers’ Keywords: Panicum virgatum, transfer guidelines, seed transfer zone, ecoregion

Abstract: Switchgrass (Panicum virgatum L.) is a perennial grass native to the North American tallgrass prairie and 

broadly adapted to the central and eastern USA. Transfer of 

germplasm throughout this region creates the potential of contaminating local gene pools with genes that are not native to 

a locale. The objective of this study was to identify structural 

patterns and spatial variation for molecular markers of switchgrass populations from the northern and central USA. Forty 

six prairie-remnant populations and 11 cultivars were analyzed 

for random amplified polymorphic DNA (RAPD) markers. 

Although there was significant population differentiation, 

little of this variation was associated with geographic regions. 

A small amount of population differentiation was associated 

with hardiness zones and ecoregions, suggesting that a recent 

proposal to use these two criteria for defining plant adaptation 

regions has merit for defining gene pools and seed-transfer 

zones of switchgrass. Cultivars of switchgrass cannot be differentiated from prairie-remnant populations in the northern 

and central USA on the basis of RAPD markers, indicating 

that they are still highly representative of natural germplasm. 

Seed sources of switchgrass can be moved considerable distance within hardiness zones and ecoregions without causing 

significant contamination, pollution, swamping, or erosion of 

local gene pools.

Link: http://afrsweb.usda.gov/SP2UserFiles/

Place/36200000/Casl-S2.pdf

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642. Darris DC, Wilson BL, Fiegener R, Johnson 

R, Horning ME. 2008. Polycross populations of the native 

grass Festuca roemeri as pre-varietal germplasm: their derivation, 

release, increase, and use. Native Plants Journal 9:305-312.

Type: Journal 

Geographic Area: Northwestern USA

Keywords: common-garden study, seed transfer zones, habitat 

restoration, erosion control, selected class release

Abstract: Results of a recent common-garden study provide 

evidence needed to delineate appropriate seed transfer zones 

for the native grass Festuca roemeri (Pavlick) E. B. Alexeev 

(Poaceae). That information has been used to develop prevariety germplasm releases to provide ecologically and genetically appropriate seeds for habitat restoration, erosion 

control, and other revegetation projects in 5 regions of the 

Pacific Northwest, U.S. Seed sources for these composite 

populations were chosen to represent a broad base of genetic 

diversity found within each region, while using plants that 

overlap in flowering time, have average to high seed yield, 

and originate at similar elevations. The process of selecting 

appropriate seed sources and developing the germplasm releases is described here. Ongoing and future investigations 

are likely to include seed production technology, establishment methods, stand management, and adaptation to diverse 

sites and specific uses.

Link: http://www.treesearch.fs.fed.us/pubs/34859

643. Davidson RH, Edwards DGW, Sziklai O, 

El-Kassaby YA. 1997. Genetic variation in germination 

parameters among populations of Pacific silver fir. Silvae Genetica 

45:2-3.

Type: Journal 

Geographic Area: Western Canada

Keywords: Abies amabilis, germination, stratification, intraclass correlation

Abstract: Strong genetic control was detected for germination capacity and germination speed in 6 populations of Pacific 

silver fir (Abies amabilis [DOUGL.] FORBES) from northern, 

central and southern Vancouver Island, British Columbia. 

Intra-class correlation estimates were high and varied between 

0.6 and 0.7 and 0.7 and 0.8 for stratified and unstratified seeds, 

respectively. Stratified seeds showed marked differences in 

germination capacity and speed among populations when compared to unstratified seeds. Family differences in germination 

determined by intra class correlation, irrespective of seed pretreatment, were the most pronounced factor when compared to 

regions and/or populations within regions. The implications of 

these differences to nursery seedling production was discussed. 

The inadvertent selection for more-rapidly germinating families in bulked seedlots may represent the most important factor 

affecting the genetic diversity of seedling crops.

largest. At the high elevation site, trees from mid- and highelevation sources grew equally well, whereas trees from 

low-elevation parents showed poorest growth. The variation 

associated with elevation zone of parent trees accounted for 

8 percent of the total experimental variation. About 9 percent 

of the total was attributed to the interaction between parent 

tree elevational zones and plantations. Height rankings of 

progenies for different ages showed little change over the 

29-year growth period of the study in low- and mid-elevation 

test sites. But at the high-elevation test site, height means for 

zone groups shifted with age as the relative heights of the 

higher elevation progenies increased.

Link: http://www.treesearch.fs.fed.us/pubs/32836

641. Czarnecki DM, Norcini JG, Zhanao D. 2007.

Phenotypic diversity of Coreopsis leavenworthii TORR. & GRAY 

(Asteraceae). Native Plants Journal 8:45-57.

Type: Journal 

Geographic Area: Florida, USA

Keywords: common-garden study, phenotypic variation, 

genetic diversity, population differentiation, wildflower

Abstract:Coreopsis leavenworthii Torr. & Gray (Asteraceae), 

one of the Florida state wildflowers, is nearly endemic to 

and very ubiquitous in Florida. Interest in commercial seed 

or plant production of this species as well as use for highway beautification, native plant community restoration, and 

mine reclamation has been increasing in recent years. This 

study aimed to understand the level and geographic distribution of phenotypic diversity of this species, a critical step 

before engaging in large-scale commercial production of 

native plants. Plants derived from natural and commercial 

seed production populations exhibited a substantial amount 

of phenotypic diversity when grown in a common garden 

study. North Florida natural populations typically had bipinnately compound leaves with dark-yellow ray flowers, while 

natural populations originating in central or south Florida 

had simpler, needle-like leaves, and light-yellow and darkyellow ray flowers. Populations were grouped into 3 clusters 

in principal component analysis. Two natural populations 

from central Florida were dissimilar and grouped into separate clusters, indicating possible existence of genetic isolation in the species’ distribution. Three production populations originating from the same native population in central 

Florida were phenotypically similar even though they had 

been grown under different climatic zones in Florida and 

represented different generations (G1, G2, or G4). This suggests that the genetic identity of the seed-source origin may 

be maintained under production practices even when plants 

are grown in locations distant from the seeds’ origin. Despite 

the clustering of these populations, there seems to be no evidence that precludes the statewide use of most populations 

in this study.

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 229

Abstract: Analysis of a common-garden study of broadleaf 

lupine (Lupinus latifolius Lindl. ex J.G.Agardh ssp. latifolius

[Fabaceae]) indicates that use of watershed delineations is better than use of plant association series for determining seed 

zones on the Mt Hood National Forest. Risk analysis further 

confirmed that only 4 seed zones are required, providing a reasonable compromise between managing costs and maintaining 

local adaptation. Overall, moderate amounts of genetic variation were found in 84 seed sources. Two principal components 

(PCs) summarized 58% of the variation in 24 measured traits, 

and variation in PC scores was significantly correlated with 

topographic, geographic, and climatic variables. Regression 

analyses showed that these variables accounted for 47% of the 

variation in the first PC and 34% of the variation in the second 

PC.

Link: http://www.nativeplantnetwork.org/Content/

Articles/6-1NPJ36-48.pdf

646. Eckert AJ, Bower AD, Gonzalez-Martinez SC, 

Wegrzyn JL, Coop G, Neale DB. 2010. Back to nature: 

ecological genomics of loblolly pine (Pinus taeda, Pinaceae).

Molecular Ecology 19:3789-3805.

Type: Journal 

Geographic Area: Southeastern USA

Keywords: adaptation, ecological genomics, environmental 

gradients, Pinus taeda, population structure, single nucleotide 

polymorphisms

Abstract: Genetic variation is often arrayed in latitudinal or 

altitudinal clines, reflecting either adaptation along environmental gradients, migratory routes, or both. For forest trees, 

climate is one of the most important drivers of adaptive phenotypic traits. Correlations of single and multilocus genotypes 

with environmental gradients have been identified for a variety 

of forest trees. These correlations are interpreted normally as 

evidence of natural selection. Here, we use a genome-wide dataset of single nucleotide polymorphisms (SNPs) typed from 

1730 loci in 682 loblolly pine (Pinus taeda L.) trees sampled 

from 54 local populations covering the full-range of the species to examine allelic correlations to five multivariate measures of climate. Applications of a Bayesian generalized linear 

mixed model, where the climate variable was a fixed effect 

and an estimated variance-covariance matrix controlled random effects due to shared population history, identified several 

well-supported SNPs associating to principal components corresponding to geography, temperature, growing degree-days, 

precipitation and aridity. Functional annotation of those genes 

with putative orthologs in Arabidopsis revealed a diverse set 

of abiotic stress response genes ranging from transmembrane 

proteins to proteins involved in sugar metabolism. Many of 

these SNPs also had large allele frequency differences among 

populations (FST = 0.10-0.35). These results illustrate a first 

step towards an ecosystem perspective of population genomics 

Link: http://www.silvaegenetica.de/fileadmin/content/dokument/archiv/silvaegenetica/45_1996/45-2-3-165.pdf

644. DeWald LE, Mahalovich MF. 2008. Historical 

and contemporary lessons from ponderosa pine genetic studies 

at the Fort Valley Experimental Forest, Arizona. In: Olberding 

SD, Moore MM, editors. Fort Valley—A Century of Research 

1908–2008. Fort Collins (CO): USDA Forest Service, Rocky 

Mountain Research Station. Proceedings RMRS-P-55. 

p 150-155.

Type: Government Document

Geographic Area: Western USA

Compilers’ Keywords: genetic integrity, Pinus ponderosa, forest management, provenance, adaptation, seed transfer, review

Abstract: Forest management will protect genetic integrity of 

tree species only if their genetic diversity is understood and 

considered in decision-making. Genetic knowledge is particularly important for species such as ponderosa pine (Pinus 

ponderosa Dougl. ex Laws.) that are distributed across wide 

geographic distances and types of climates. A ponderosa pine 

study initiated in 1910 at the Fort Valley Experimental Forest 

is among the earliest ponderosa pine genetic research efforts 

in the United States. This study contributed to the description 

of ponderosa pine’s varietal differences, genetic diversity and 

adaptation patterns, and helped confirm the importance of using local seed sources. The role this and other pioneer studies 

had in improving forest management of ponderosa pine was, 

and still is critical. These early studies have long-term value 

because they improve our knowledge of responses to climate 

change and our understanding of genetic variability in physiology and pest resistance in older trees. More recently, studies 

of natural ponderosa pine stands at Fort Valley using molecular markers have shown the importance of stand structure and 

disturbance regimes to genetic composition and structural 

patterns. This knowledge is important to ensure ecological 

restoration efforts in ponderosa pine forests will also restore 

and protect genetic integrity into the future. Highlights of these 

historical and contemporary studies at Fort Valley are summarized and their applications to management of ponderosa pine 

forests are described.

Link: http://www.treesearch.fs.fed.us/pubs/32505

645. Doede DL. 2005. Genetic variation in broadleaf lupine 

(Lupinus latifolius) on the Mt Hood National Forest and implications for seed collection and deployment. Native Plants Journal 

6:36-48.

Type: Journal 

Geographic Area: Oregon, USA

Keywords: adaptation, seed transfer risk, seed transfer zones, 

common-garden

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230 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Abstract: Adaptation to precipitation conditions may induce 

genetic diversity that changes morphological and physiological traits. This hypothesis was investigated in the seedlings 

of seven western redcedar (Thuja plicata Donn ex D. Don) 

populations, which were collected along a precipitation transect from the Pacific coast to the southern interior of British 

Columbia, Canada. The experimental seedlings were either 

well-watered or soil-droughted and measured for growth, gas 

exchange rates, transpiration efficiency, and carbon isotope 

discrimination during or at the end of the third growing season. Significant variation was found in most of these morphological and physiological traits among the populations. Much 

of this variation occurred under well watered, but not so much 

under droughted conditions. Mean height increments and 

transpiration efficiency showed a significant linear relationship, but biomass increments exhibited a quadratic relationship with precipitation on the origin site of these populations. 

Measurements of water use efficiency obtained from instantaneous gas exchange measurements, carbon isotope discrimination, and transpiration efficiency were intercorrelated in the 

seedlings. However, neither did any of these measurements 

consistently rank the populations, nor were they indicative of 

adaptation to climatic precipitation conditions in these western redcedar populations.

Link: http://www.researchgate.net/publication/216860712_

Morphological_and_physiological_variation_in_western_redcedar_(Thuja_plicata_Donn._Ex_D._Don)_populations_under_contrasting_soil_water_conditions/

file/79e4150f5b1e8c69c6.pdf

649. Fant JB, Holmstrom RM, Sirkin E, Etterson 

JR, Masi S. 2008. Genetic structure of threatened native 

populations and propagules used for restoration in a clonal 

species, American beachgrass (Ammophila breviligulata Fern.).

Restoration Ecology 16:594-603.

Type: Journal 

Geographic Area: Central USA

Keywords: American beachgrass (Ammophila breviligulata 

Fern.), clonal plants, ISSR, molecular diversity, population genetic structure, restoration genetics

Abstract: An important goal of native plant restorations 

was to reconstitute populations that are genetically similar 

to native ones, thereby increasing the probably of successful establishment and persistence. We examined the extent 

to which this goal has been accomplished in Great Lakes 

restorations of Ammophila breviligulata Fern., a beachgrass species that is widely used for habitat restoration 

and is considered threatened in the study areas. In parallel studies on Lake Michigan and Lake Superior, we used 

polymorphic Intersimple Sequence Repeat markers to assess genetic similarity between well-established and new 

native populations, restored populations, and restoration 

for non-model organisms, but also highlight the need for further integration of the methodologies employed in spatial statistics, population genetics and climate modeling during scans 

for signatures of natural selection from genomic data.

Link: http://www.ncbi.nlm.nih.gov/pubmed/20723060

647. Erickson VJ, Mandel NL, Sorensen FC. 2004.

Landscape patterns of phenotypic variation and population 

structuring in a selfing grass, Elymus glaucus (blue wildrye).

Canadian Journal of Botany 82:1776-1789.

Type: Journal 

Geographic Area: Oregon, USA

Keywords: Elymus glaucus, morphological variation, local adaptation, seed transfer, seed zones, polyploidy

Abstract: Source-related phenotypic variance was investigated in a common garden study of populations of Elymus 

glaucus Buckley (blue wildrye) from the Blue Mountain 

Ecological Province of northeastern Oregon and adjoining 

Washington. The primary objective of this study was to assess 

geographic patterns of potentially adaptive differentiation in 

this self-fertile allotetraploid grass, and use this information 

to develop a framework for guiding seed movement and 

preserving adaptive patterns of genetic variation in ongoing 

restoration work. Progeny of 188 families were grown for 3 

years under two moisture treatments and measured for a wide 

range of traits involving growth, morphology, fecundity, and 

phenology. Variation among seed sources was analyzed in 

relation to physiographic and climatic trends, and to various 

spatial stratifications such as ecoregions, watersheds, edaphic classifications, etc. Principal component (PC) analysis 

extracted four primary PCs that together accounted for 67% 

of the variance in measured traits. Regression and cluster 

analyses revealed predominantly ecotypic or stepped-clinal 

distribution of genetic variation. Three distinct geographic 

groups of locations accounted for over 84% of the variation 

in PC-1 and PC-2 scores; group differences were best described by longitude and ecoregion. Clinal variation in PC-3 

and PC-4 scores was present in the largest geographic group. 

Four geographic subdivisions were proposed for delimiting

E. glaucus seed transfer in the Blue Mountains.

Link: http://www.fs.fed.us/pnw/pubs/journals/pnw_2004_

erickson001.pdf

648. Fan S, Grossnickle SC, Russell JH. 2008.

Morphological and physiological variation in western redcedar 

(Thuja plicata) populations under contrasting soil water 

conditions. Trees 22:671-683.

Type: Journal 

Geographic Area: Western North America

Keywords: carbon isotope discrimination, gas exchange, precipitation, adaptation, water use efficiency

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 231

study investigates the potential trade-off between seedling 

growth and alkaloid production, and associations between 

topographic/climatic variables and alkaloid production. 

Piperidine alkaloids were quantified in foliage of 501 

nursery seedlings grown from seed sources in west-central Washington, Oregon and California, roughly covering 

the western half of the native range of ponderosa pine. A 

nested mixed model was used to test differences among 

broad-scale regions and among families within regions. 

Alkaloid concentrations were regressed on seedling 

growth measurements to test metabolite allocation theory. 

Likewise, climate characteristics at the seed sources were 

also considered as explanatory variables. Quantitative 

variation from seedling to seedling was high, and regional variation exceeded variation among families. Regions 

along the western margin of the species range exhibited 

the highest alkaloid concentrations, while those further 

east had relatively low alkaloid levels. Qualitative variation in alkaloid profiles was low. All measures of seedling 

growth related negatively to alkaloid concentrations on 

a natural log scale; however, coefficients of determination were low. At best, annual height increment explained 

19.4% of the variation in ln (total alkaloids). Among the 

climate variables, temperature range showed a negative, 

linear association that explained 41.8% of the variation. 

Given the wide geographic scope of the seed sources and 

the uniformity of resources in the seedlings’ environment, observed differences in alkaloid concentrations are 

evidence for genetic regulation of alkaloid secondary metabolism in ponderosa pine. The theoretical trade-off with 

seedling growth appeared to be real, however slight. The 

climate variables provided little evidence for adaptive alkaloid variation, especially within regions.

Link: http://www.ncbi.nlm.nih.gov/pubmed/19010800

652. Gibbs JP, Smart LB, Newhouse AE, Leopold 

DJ. 2012. A molecular and fitness evaluation of commercially 

available versus locally collected blue lupine Lupinus perennis

L. seeds for use in ecosystem restoration efforts. Restoration 

Ecology 20:456-461.

Type: Journal 

Geographic Area: New York, USA

Keywords: blue lupine, genetics, habitat restoration, Karner 

blue butterfly, Lupinus perennis, Lycaeides melissa samuelis, microsatellite, seed source

Abstract: Dependence on wild seed sources is often impractical for large-scale habitat restoration programs. Reliance on 

commercial seed supplies of unknown provenance and fitness is 

thereby warranted. Little consideration has been given, however, 

to how the large volumes of seed required should be sourced. We 

evaluated commercial and locally collected seed sources for potential use in a New York State-based, landscape-scale program 

propagules obtained from two commercial suppliers. 

Native populations were generally more diverse than 

expected for a clonal species, whereas the commercially 

cultivated releases were monotypic. One of the commercial releases used in Minnesota was exclusively found in 

restored populations and did not occur in any other native 

population at this site. The propagules used in the newly 

planted restoration in Illinois were derived from a release 

that commercial suppliers maintain was derived from a native Michigan population, as opposed to a selected release. 

Diversity in this restoration was equivalent to that native 

Illinois’ populations; however, many of the genotypes 

were not of local origin. Overall, study underscores the 

importance of obtaining baseline genetic surveys of remnant native populations and restoration propagules before 

restoration efforts are initiated, especially when the populations are threatened or endangered.

650. Ferrell WK, Woodward ES. 1966. Effects of 

seed origin on drought resistance of Douglas-fir (Pseudotsuga 

menziesii) (Mirb.) Franco. Ecology 47:499-503.

Type: Journal 

Geographic Area: Pacific Northwest, USA, Canada

Compilers’ Keywords: drought hardiness, population 

variation

Abstract: Douglas-fir seedlings from a number of seed origins 

were subjected to drought conditions in the greenhouse and 

laboratory. Interior mountain seedlings showed significantly 

greater drought resistance than seedlings from origins west of 

the Cascade Mountains. Differences were also found within 

each of these groupings. In the Corvallis, Oregon, area seedlings 

produced from seed on a south slope had more drought resistance than those from a short distance away on a north slope. 

Differences in drought resistance may involve either drought 

hardiness or drought avoidance, or both.

651. Gerson EA, Kelsey RG, St. Clair JB. 2009.Genetic 

variation of piperidine alkaloids in Pinus ponderosa: a common 

garden study. Annals of Botany 103:447-457.

Type: Journal 

Geographic Area: Western USA

Keywords: Pinus ponderosa var. ponderosa, Pinacea, 2,6-disubstituted piperidine alkaloids, secondary products, geographic 

variation, progeny study, plant defense, Growth-Differentiation 

Balance Hypothesis, PRISM

Abstract: Previous measurements of conifer alkaloids 

have revealed significant variation attributable to many 

sources, environmental and genetic. The present study 

takes a complementary and intensive, common garden 

approach to examine genetic variation in Pinus ponderosa var. ponderosa alkaloid production. Additionally, this 

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232 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

species, and other species in the family Pinaceae. No significant difference was detected among stands in outcrossing rate. 

The high level of inbreeding observed in this population could 

be attributable to population fragmentation due to human activities, or to reduced levels of pollen production in this population compared to other populations of ponderosa pine. Genetic 

conservation strategies for this population should include the 

protection of the large native stands that remain and the use of 

local, adapted sources for afforestation. Management of pollen 

in a seed orchard should improve the genetic quality of seed 

over wild stand seed by reducing inbreeding levels.

Link: http://scholarsarchive.library.oregonstate.edu/xmlui/

handle/1957/10704

654. Gordon DR, Rice KJ. 1998. Patterns of differentiation 

in wiregrass (Aristida beyrichiana): implications for restoration 

efforts. Restoration Ecology 6:166-174.

Type: Journal 

Geographic Area: Florida, USA

Compilers’ Keywords: local adaptation, genetic diversity, 

longleaf pine, common garden, reciprocal transplant

Abstract: Aristida beyrichiana (wiregrass) is increasingly 

being planted in restoration projects across the southeastern 

coastal plain, with little focus on genetic differences among 

populations across the region. Local and regional population 

differentiation for establishment and growth traits were examined in common garden and reciprocal transplant experiments. Seeds from up to 20 plants from each of seven populations were collected in northern and central Florida sites that 

encompassed gradients of soils, hydrology, and temperature. 

Reciprocal seed transplants using three of the common garden 

populations were conducted in two consecutive years. In the 

common garden, significant population differences were seen 

in seed weight, seedling emergence and survival, tiller height, 

number of tillers, the relationship between tiller number and 

tiller height, and flowering. Variation among maternal families 

was seen in tiller number and in the relationship between tiller 

number and tiller height. The reciprocal transplant study did 

not detect either local adaptation to sites of origin or consistent 

superiority of one source population or planting site in seedling establishment. These results suggest that the probability of 

seedling establishment is primarily dependent on environmental conditions rather than genetic differences. Genetic variation 

for traits related to fitness (e.g., tiller number) may be retained 

within populations because phenotypically plastic growth responses of seedlings to environmental variation buffer genetic 

variation against the action of selection. But despite the lack 

of evidence for genetic influences on initial establishment in 

wiregrass, our common garden study suggests genetic differences among populations. This result, when combined with 

previous results indicating local adaptation in later life stages 

of wiregrass, suggests that restoration efforts involving this 

for restoring blue lupine Lupinus perennis. Through analysis of 

microsatellite markers we determined that “native” lupine designations by some commercial suppliers were in fact interspecific hybrids and therefore unreliable; at least two commercial 

sources, however, were genetically as close to native New York 

populations as native New York populations were to one other. 

Common garden experiments revealed that seed source influenced first-year overwintering survival and subsequent height 

growth of surviving plants; seed sources more closely related 

genetically to native New York populations survived better and 

produced more stems per individual in the field in the area targeted for restoration. We conclude that (1) commercial suppliers 

often but not always offer reliably characterized seed sources of 

sufficient genetic similarity to native populations to warrant their 

use in restoration projects and (2) genetic affinity of potential 

seed stock to native populations is positively related to its fitness 

in the environment targeted for restoration.

653. Gooding GD. 1998. Genetic variation and mating 

system of ponderosa pine in the Willamette Valley of Oregon 

[Thesis]. Corvallis (OR): Oregon State University. 104 p.

Type: Thesis

Geographic Area: Pacific Northwest USA

Compilers’ Keywords: genetic structure, genetic diversity, 

Pinus ponderosa, outcrossing, inbreeding and outbreeding depression, fragmentation, pollen management

Abstract: The population genetic structure of ponderosa pine 

(Pinus ponderosa Dougl.) in the Willamette Valley of Oregon 

was investigated. Cones were collected from native stands of 

ponderosa pine from the Willamette Valley, eastern Oregon, 

southwest Oregon and the Puget Sound Basin of Washington. 

Seeds were subjected to isozyme analysis of 12 enzyme systems controlled by 20 different loci. Population genetic parameters for progeny, parent trees and pollen were estimated from 

isozyme data using BIOSYS-2, a population genetic statistical software package. For progeny data, total gene diversity 

(HT) was 0.249; average genic diversity within stands (Hs) was 

0.231; average genic diversity among stands (DST) was 0.017; 

the proportion of total genic diversity due to differences among 

stands (GST) was 0.067. Based on gene diversity statistics and 

genetic distances, stands of Willamette Valley ponderosa pine 

do not differ substantially from one another or from other populations of ponderosa pine in Washington and Oregon. These 

results suggest that protection of the Willamette Valley population can be accomplished by conserving a large number of individuals in a few large stands or many small stands. Multilocus 

and single-locus estimates of outcrossing rate were determined 

for six stands within the Willamette Valley. Outcrossing rates 

were estimated for seven polymorphic loci using MLTR, a 

mating system analysis software package. Estimates of multilocus outcrossing for these stands (mean 0.642; range 0.365 

to 0.879) were substantially lower than estimates previously 

reported for other populations of ponderosa pine, other pine 

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 233

first overview about possible effects. The analysis was then 

extended using Accelerated Failure Time (AFT) models to estimate the parameters of a Weibull function, used to describe 

survival times. The resulting models were used to simultaneously analyze the effects of intensity of mixture (represented 

by Shannon–Weaver-Index and, alternatively, by species proportion), time since harvest and site characteristics. Results obtained indicate positive effects of species mixture on resistance 

of spruce trees: survival probabilities increase with increasing 

intensity of mixture, regardless whether mixture is characterized by Shannon–Weaver-Index or species proportion. Spruce 

trees in monocultures on average site conditions will reach 

age 100 with a probability of 80%. Spruce trees growing in a 

moderately mixed stand(average Shannon–Weaver-Index 0.4) 

show a slight increase in survival probability to a 83% probability of reaching age 100 whilst spruce trees in a more diverse 

stand (average Shannon–Weaver-Index 1.2) have a 97% probability of reaching age 100. An admixture of 50% thus leads to 

an increase in survival probability of 17 percentage points. Site 

variables even show a stronger impact on survival than tree 

species mixture. From these variables wet soils had the strongest negative influence on spruce survival, while orographic 

conditions of saddles, anticlines, valleys, trenches or dells 

showed the strongest positive influence on survival. However, 

the strongest influence on spruce survival was recent harvest 

activity. The more time had passed since the harvest operation, 

the less likely residual trees were to succumb to stresses.

[Note: Griess VC, Acevedo R, Härtl F, Staupendahl K, 

Knoke T. 2012. Erratum to “Does mixing tree species enhance 

stand resistance against natural hazards? A case study for spruce”

[Forest Ecol. Manage. 267 (2012) 284–296]. Forest Ecology 

and Management 276:259.]

657. Griffin AR, Ching KK. 1977. Geographic variation in Douglas-fir from the coastal ranges of California. Silvae 

Genetica 26:149-157.

Type: Journal 

Geographic Area: California, USA

Compilers’ Keywords: Pseudotsuga menziesii, common garden, bud set, cold hardiness

Summary: 181 wind-pollinated families of Douglas-fir 

from through-out the coastal ranges of northern California 

were raised in a nursery at Corvallis, Oregon and assessed 

for growth, phenology, cold hardiness, and response to moisture stress. Variation patterns for seed weight, germination 

rate and cotyledon number were also determined. All characters varied genetically, and with the exception of date of 

bud burst the most significant contrast was between samples 

from the coastal fog-belt and those from the interior ranges. Coastal seed was relatively small and germinated more 

slowly, and seedlings had fewer cotyledons, less hypocotyl, 

species should use local seed sources from sites with similar 

soil and hydrological conditions.

655. Goto S, Iijima H, Ogawa H, Ohya K. 2011.

Outbreeding depression caused by intraspecific hybridization 

between local and nonlocal genotypes in Abies sachalinensis.

Restoration Ecology 19:243-250.

Type: Journal 

Geographic Area: Japan

Keywords: Abies sachalinensis, ecological restoration, elevation, intraspecific hybridization, local adaptation, outbreeding 

depression

Abstract: To evaluate the effects of intraspecific hybridization of local and nonlocal genotypes on growth traits of progeny in Abies sachalinensis, we performed reciprocal crossing 

between nonlocal trees in a high-elevation zone (1,100–1,200 

m asl) and local trees in a low-elevation zone (530 m asl) in 

Hokkaido, northern Japan, in 1979 and established a common 

garden experiment using local × local (female × male), local × 

nonlocal, nonlocal × local, and nonlocal × nonlocal progeny in 

the low-elevation zone in 1986. Survival, height, diameter at 

breast height (dbh), needle nitrogen content, specific leaf area 

(SLA), and needle area per shoot diameter (NA) of 25-year-old 

progeny were measured in 2005. The survival rate was consistently high (> 85% on average). Reductions in height and 

dbh were apparent in F1 hybrids compared with local × local 

progeny. Furthermore, outbreeding depression was significant 

in height growth of nonlocal × local F1 hybrids and in dbh of 

both F1 hybrids. Reductions in growth traits may be related to 

morphological needle traits, such as the low values of SLA and 

NA. Elevation guidelines for A. sachalinensis seed zones are 

discussed to ensure the long-term viability of both restored and 

native populations.

656. Griess VC, Acevedo R, Härtl F, Staupendahl 

K, Knoke T. 2012. Does mixing tree species enhance stand 

resistance against natural hazards? A case study for spruce. Forest 

Ecology and Management 267:284-296.

Type: Journal 

Geographic Area: Germany, Europe 

Keywords: mixed species stand, survival analysis, risk, forest 

management, forest damage survey, Weibull

Abstract: In this study, survival of spruce (Picea abies [L.] 

Karst.) trees in mixed- and mono-species stands was analyzed 

using the database of Rhineland Palatinate’s forest damage 

survey (FDS). The influence of species mixture on tree survival probability was analyzed using data from 9,864 trees, of 

which 2,866 spruce trees have been analysed in detail. Data 

was collected on 495 research plots in a series of continuous 

measurements taken since 1984. For estimating survival probability, the Kaplan–Meier method was applied to achieve a 

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234 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Abstract: Variation in the ability of western redcedar (Thuja 

plicata Donn ex D. Don) populations to withstand water stress 

may exist because this species is found in coastal and interior 

biogeoclimatic subzones representing the full range of precipitation regimes in British Columbia, Canada. Seven western redcedar populations from locations in British Columbia, 

representing a wide range of habitat types, were assessed for 

their gas exchange and water relations response to controlled 

drought. Before drought, population variation occurred in stomatal conductance, net CO2 assimilation rate and intrinsic water use efficiency and the relative water content at turgor loss 

point. During drought, populations had different responses of 

net CO2 assimilation to decreasing predawn shoot water potential. After drought, populations differed in stomatal conductance and intrinsic water use efficiency, plus osmotic potential 

at turgor loss point, osmotic potential at saturation and apparent cuticular transpiration. Western redcedar populations from 

drier-inland habitats had a lower osmotic potential at turgor 

loss point, lower relative water content at turgor loss point and 

lower apparent cuticular transpiration in response to drought 

than populations from coastal origin with temperate maritime 

habitat. Reduction of cuticular water loss and adjustments of 

cellular water relations in response to drought was found to 

occur among seven western redcedar populations originating 

along a precipitation gradient while; there were minimal population differences in the gas exchange response to drought.

Link: http://www.researchgate.net/publication/216860649_

Physiological_variation_among_western_redcedar_(Thuja_

plicata_Donn._Ex_D._Don)_populations_in_response_to_

short-term_drought/file/9fcfd50f5b1b7e68c0.pdf

660. Grossnickle SC, Sutton BCS, Folk RS, Gawley 

RJ. 1996. Relationship between nuclear DNA markers and 

physiological parameters in Sitka x interior spruce populations. 

Tree Physiology 16:547-555.

Type: Journal 

Geographic Area: Western Canada

Keywords: freezing tolerance, nuclear DNA markers, Picea 

engelmannii, Picea glauca, Picea sitchensis, water relations 

parameters

Summary: Eight populations of Sitka spruce (Picea sitchensis (Bong.) Carr.) and interior spruce (Picea glauca (Moench) 

Voss × Picea engelmannii Parry ex. Engelm.) seedlings were 

sampled from a zone of Sitka-interior spruce introgression in 

British Columbia, Canada. Restriction fragment length polymorphisms of the nuclear ribosomal RNA genes (rDNA) were 

used to define species-specific hybridization patterns for the 

Sitka spruce and interior spruce populations. Hybridization 

was estimated from an index based on the relative abundance 

of polymorphic rDNA combining bands for each population. 

Sitka × interior hybrid seedlings had an index value for the relative abundance of interior spruce rDNA (Si-rDNA) ranging 

but greater epicotyl growth; grew for a longer period before 

setting buds; showed less capacity to set buds in response to 

moisture stress; and were less cold hardy. Lesser variation 

patterns were associated with elevation and latitude. In spite 

of broad similarities in variation patterns, the distribution of 

variation among sampling levels was not the same for all 

characters. For example, time of bud set and epicotyl growth 

were closely associated with the regional difference in climate, the ratio of geographic : within stand variation being 

> 10, compared with values < 1 for seed-related characters. 

Cold hardiness variation was also predominantly regional. 

Such differences indicate a variable complex of selection 

pressure gradients, and militate against any systematic distinction of the coastal and interior range populations. 

Link: http://silvaegenetica.com/fileadmin/content/dokument/archiv/silvaegenetica/26_1977/26-5-6-149.pdf

658. Grossnickle SC, Sutton BCS. 1999. Applications 

of biotechnology for forest regeneration. New Forests 

17:213-226.

Type: Journal 

Geographic Area: Canada

Keywords: tissue culture, molecular genetics, microbial inoculations, ecophysiology, Picea, somatic embryo

Abstract: The Forest Biotechnology Centre is an interdisciplinary research group dedicated to the development and application of advanced technology for the enhancement of forest regeneration. The Centre carries out contracts on behalf of 

clients in forest-related industries and government agencies. In 

addition, there are a number of long-term, in-house projects 

aimed at the development of proprietary technologies in genetics and propagation, and seedling production and establishment. Technical capabilities include: tissue culture, molecular 

genetics, pathology and microbial inoculants, and ecophysiology. These techniques are also being used to improve nursery 

culture regimes, disease assessment, planting regimes, and 

new product development for a variety of conifer species. 

Additional programs relate population genetics to adaptive 

traits, and develop clonal testing within elite families from 

tree-breeding programs.

Link: http://www.cbd.int/doc/articles/2008/A-00665.pdf

659. Grossnickle SC, Russell JH. 2010. Physiological 

variation among western redcedar (Thuja plicata Donn ex D. Don) 

populations in response to short-term drought. Annals of Forest 

Science 67:506-506.

Type: Journal 

Geographic Area: Western Canada

Keywords: western redcedar population, performance, gas exchange, water relations, drought response

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 235

Compilers’ Keywords: genetic variation, genecology, climate 

change, seed transfer guidelines, tree improvement

Abstract: In this thesis genecology, phylogeography, and 

quantitative genetics of red alder were investigated. Further, 

literature with respect to red alder product values, market demand, forest resources, and red alder biology were reviewed. 

Implications of these aspects for genetic resources management 

were evaluated and a breeding strategy for red alder in British 

Columbia was developed. Genetic differentiation among 

provenances in British Columbia was investigated based on 

six polymorphic allozyme loci and measurements of six quantitative traits. Multivariate analysis revealed complex associations of quantitative traits with the latitude, distance to the 

coast, and elevation of the seed source. Nei’s genetic distance 

revealed a strong differentiation among island and mainland 

provenances at one allozyme locus. This differentiation can be 

interpreted as a result of migration from two different refugia 

since the last glaciation. Adaptation of red alder provenances 

was investigated based on trials in multiple planting environments. Significant genotype × environment interactions were 

found at the population and family level. Provenances close to 

each planting site showed superior performance in growth and 

survival, suggesting adaptation of red alder to local environments. Seed transfer guidelines and seed procurement zones 

were developed under the assumption of local optimality using improved methodology based on risk associated with seed 

transfer. In order to assess the potential benefits from selection 

and evaluate different options for tree improvement, genetic 

parameters for growth and adaptive traits were estimated from 

progeny tests. Estimated heritabilities for growth and adaptive traits were moderate with values between 0.30 and 0.50. 

Genetic gain in growth traits from individual selection would 

range from 25 to 35%. Small improvements (approximately 

5%) would indirectly be achieved in the form score due to 

positive genetic correlations. Removal of spatially autocorrelated error variation in field experiments increased heritabilities and gains from selection. A multiple population breeding 

strategy for uncertain climatic conditions in British Columbia 

was derived as a synthesis, using information on genecology, 

risk associated with seed transfer, and genetic parameters. Six 

breeding populations of red alder, some with novel trait combinations, were found to be necessary to cope with possible 

effects of climatic warming in British Columbia.

Link: http://www.ualberta.ca/~ahamann/publications/pdfs/

Hamann_1999_Abstract.pdf

663. Hamann A, Koshy MP, Namkoong G, Ying 

CC. 2000. Genotype x environment interactions in Alnus rubra: 

developing seed zones and seed-transfer guidelines with spatial 

statistics and GIS. Forest Ecology and Management 136:107-119.

Type: Journal 

Geographic Area: Western Canada

from 0.07 (Lower Nass; the most westerly collected source) 

to 0.95 (Bulkley Valley low-elevation seed orchard). During 

shoot elongation, osmotic potential at saturation (Ψsat) and turgor loss point (Ψtlp) increased, whereas total turgor (ΨPTotal) 

decreased. After bud set in the summer and throughout the 

fall, Ψsat and Ψtlp decreased, whereas ΨPTotal increased. At all 

times of year, populations with a higher Si-rDNA index had 

lower Ψtlp and Ψsat and higher ΨPTotal than populations with 

a lower Si-rDNA index. During the fall, Sitka × interior hybrid seedlings exhibited a seasonal decline in the temperature 

causing 50% needle electrolyte leakage (LT50) and in the critical temperature indicating the initial point of freezing injury. 

Seedlings with a higher Si-rDNA index had lower LT50 and 

critical temperature values indicating greater freezing tolerance in the fall. Throughout most of the year, seedling population Si rDNA index was related to the degree of drought and 

freezing tolerance.

661. Gylander T. 2011. The potential of aspen clones and 

hybrids for enhanced forest management in Alberta [Thesis].

Edmonton, Alberta, Cananda: University of Alberta. 112 p.

Type: Thesis

Geographic Area: Western Canada

Compilers’ Keywords: aspen management, Populus 

tremuloides, genetic variation, assisted migration, provenance, forest productivity

Abstract: This thesis presents results from an industrial aspen 

tree improvement program for Alberta, evaluating a series of 

provenance, clonal and hybrid field trials. The goals were to 

(1) investigate geographic patterns of genetic variation in order to delineate breeding regions, (2) to assess the potential of 

clonal forestry systems to enhance forest productivity, and (3) 

to evaluate the potential of hybridization to enhance growth 

through hybrid vigor. Partitioning of genetic variance with geographic predictor variables suggests two breeding regions for 

Alberta should be appropriate: a Sub-Boreal Rocky Mountain 

Foothill region between 52°30’ N and 56° N latitude, and a 

Boreal Mixedwood region between 56° N and 59° N latitude. 

Broad-sense heritabilities for height and diameter ranged from 

0.36 to 0.64 on selected sites, allowing 5–15% genetic gains in 

height and 9–34% in diameter based on selections from current 

trials. The best genotypes within hybrid families could have 

some additional potential in improving yields.

Link: http://www.ualberta.ca/~ahamann/people/pdfs/Gylander_2011_MSc.pdf

662. Hamann A. 1999. Utilization and management of 

red alder genetic resources in British Columbia [Dissertation]. 

Vancouver, British Columbia, Canada: The University of British 

Columbia. 13 p.

Type: Dissertation

Geographic Area: Western Canada

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236 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Abstract: A three year common garden study was conducted 

on whitebark pine (Pinus albicaulis) which included 215 families from the eight provenances or seed zones in Oregon and 

Washington. Total height and needle color were assessed. 

Height differed significantly among provenances and families, 

and was primarily associated with source elevation, longitude, 

and precipitation. A moderate to high heritability was estimated for total height. Seedling needle color differed among 

provenances and was associated with temperature : moisture indexes and source elevation. Height growth along the 

Cascade mountain range (USA), representing four adjoining 

provenances or seed zones, appeared to be similar and clinal 

in nature. The four remaining provenances, representing seed 

zones from a more moderate or extreme environmental settings, differed significantly from the Cascade provenances for 

one or more traits that were examined. It would seem prudent 

to restrict seed transfers to within each of these four seed zones 

respectively. This study also supports the need to plan gene 

conservation collections within each seed zone for whitebark 

pine in the Pacific Northwest region.

Link: http://www.treesearch.fs.fed.us/pubs/38209

665. Hancock N, Leishman MR, Hughes L. 2012.

Testing the “Local Provenance” paradigm: a common garden 

experiment in Cumberland Plain Woodland, Sydney, Australia. 

Restoration Ecology 21:569-577.

Type: Journal 

Geographic Area: Australia

Keywords: adaptive potential, home-site advantage, local adaptation, local superiority, restoration, seed source

Abstract: Seed for restoration projects has traditionally 

been sourced locally to “preserve” the genetic integrity of 

the replanted site. Plants grown from locally sourced seeds 

are perceived to have the advantage of being adapted to local conditions, and the use of local provenance is a requirement of many restoration projects. However, the processes 

of climate change and habitat fragmentation, with the subsequent development of novel environments, are forcing us to 

reconsider this basic tenet of restoration ecology. We tested 

the “local provenance is best” paradigm, by comparing the 

performance of plants grown from local with non-local seed 

sources within a common garden experiment. We selected six 

species representing a range of growth forms (Acacia falcata, 

Bursaria spinosa ssp. spinosa, Eucalyptus crebra, E. tereticornis, Hardenbergia violacea and Themeda australis) from an 

assemblage known as the Cumberland Plain Woodland, a 

threatened community in western Sydney. Multiple provenances were collected from within the range of each species and grown at two field sites on the Cumberland Plain. 

Growing time varied between species and ranged from 7 

months to 2 years. With the exception of B. spinosa, and to a 

lesser extent T. australis, we found little evidence that local 

Keywords: red alder, genecology, seed zones, seed transfer 

guidelines, Kriging, British Columbia

Abstract: Multiple provenance trials of red alder (Alnus 

rubra Bong.) were used to demonstrate how geostatistical methods can be applied to develop better seed-transfer 

guidelines and seed-procurement zones for forestry. Height 

and survival of 65 provenances from British Columbia were 

measured at four test sites. Significant genotype-environment interactions were found at the population and family 

level. Provenances close to each planting site showed superior performance in growth and survival, suggesting adaptation of red alder to local environments. The environmental 

basis of local adaptations was evaluated using redundancy 

analysis. Geographic and climatic variates accounted for 

approximately equal amounts of the variation in height 

growth (21 and 23%, respectively), while geographic variates accounted for 60% of the variation in climatic variables. 

Loadings of the redundancy variates suggested that both, 

gene flow and selection by environmental factors shaped 

geographic patterns of genetic differentiation in red alder. 

Performance of seed sources at unknown locations was predicted with ordinary kriging throughout the natural range of 

red alder in British Columbia. General seed-transfer guidelines were then generated with principal component analysis 

of predicted reaction norms. We found clinal differentiation 

of reaction norms along the coast from northwest to southeast. Further, the Georgia depression was identified as an 

area of genetic differentiation. Based on predicted height 

and survival at two trials that most realistically represented 

operational planting sites, we derived simple rules for seed 

transfer. A transfer of 100 km in either direction along the 

coast was associated with a decline of 2.5% in survival, 

and 5 cm in 2-year height. Finally, we showed how maps 

of predicted performance and associated variance surfaces 

can be used to develop seed zones for specific objectives, 

where seed zones are given as maps of probabilities of a seed 

source performing above or below a given threshold for any 

combination of traits.

Link: http://www.ualberta.ca/~ahamann/publications/pdfs/

Hamann_et_al_2000.pdf

664. Hamlin J, Kegley A, Sniezko R. 2011. Genetic 

variation of whitebark pine (Pinus albicaulis) provenances and 

families from Oregon and Washington in juvenile height growth 

and needle color. In: Keane RE, Tomback DF, Murray MP, Smith 

CM, editors. High Five Symposium. Fort Collins (CO): USDA 

Forest Service, Rocky Mountain Research Station. Proceedings 

RMRS-P-63. p 133-139.

Type: Government Document

Geographic Area: Pacific Northwest USA

Compilers’ Keywords: common garden, seed zones, gene 

conservation, seed collection

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 237

allozymes. Therefore, these traits appear to be under strong 

natural selection. Nonetheless, high levels of genetic variation 

persist within populations. The genetic control of cold adaptation traits ranges from weak to strong, with phenological traits 

having the highest heritabilities. Within-population genetic 

correlations among traits range from negligible to moderate. 

Generally, bud phenology and cold hardiness in the fall are genetically uncorrelated with bud phenology and cold hardiness 

in the spring. Analyses of quantitative trait loci indicate that 

cold adaptation traits are mostly controlled by multiple genes 

with small effects and that quantitative trait loci × environment 

interactions are common. Given this inherent complexity, we 

suggest that future research should focus on identifying and 

developing markers for cold adaptation candidate genes, then 

using multilocus, multiallelic analytical techniques to uncover 

the relationships between genotype and phenotype at both the 

individual and population levels. Ultimately, these methods 

may be useful for predicting the performance of genotypes in 

breeding programs and for better understanding the evolutionary ecology of forest trees.

Link: http://www.treesearch.fs.fed.us/pubs/42203

668. Hufford KM, Mazer SJ. 2003. Plant ecotypes: 

genetic differentiation in the age of ecological restoration. Trends 

in Ecology and Evolution 18:147-155.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: local adaptation, seed transfer, translocation, introduced, genetic dilution, restoration genetics, review, 

seed zones, common garden

Abstract: Recent studies illustrate the emerging field of restoration genetics, which is a synthesis of restoration ecology 

and population genetics. The translocation of organisms during the restoration of native ecosystems has provoked new 

questions concerning the consequences of sampling protocols 

and of intraspecific hybridization between locally adapted and 

transplanted genotypes. Studies are now underway to determine both the extent of local adaptation among focal populations and the potential risks of introducing foreign genotypes, 

including founder effects, genetic swamping and outbreeding 

depression. Data are needed to delineate ‘seed transfer zones’, 

or regions within which plants can be moved with little or no 

consequences for population fitness. Here, we address the 

revival of transplant and common garden studies, the use of 

novel molecular markers to predict population genetic consequences of translocation, and their combined power for determining appropriate seed transfer zones in restoration planning 

for native plant populations.

Link: http://www.researchgate.net/publication/222650495_

Plant_ecotypes_genetic_differentiation_in_the_age_of_ecological_restoration/file/60b7d517ead892c10a.pdf

provenance plants were superior to distant provenances in 

terms of survival and establishment.

666. Horning ME, McGovern TR, Darris DC, 

Mandel NL, Johnson RC. 2010.Genecology of Holodiscus 

discolor (Rosaceae) in the Pacific Northwest, U.S.A. Restoration 

Ecology 18:235-243.

Type: Journal 

Geographic Area: Western USA

Keywords: common garden, genecology, germplasm, 

Holodiscus discolor, oceanspray

Abstract: An important goal for land managers is the incorporation of appropriate (e.g., locally adapted and genetically diverse) plant materials in restoration and revegetation 

activities. To identify these materials, researchers need to 

characterize the variability in essential traits in natural populations and determine how they are related to environmental 

conditions. This common garden study was implemented to 

characterize the variability in growth and phenological traits 

relative to climatic and geographic variables of 39 Holodiscus 

discolor (Pursh) Maxim. accessions from locations throughout 

the Pacific Northwest, U.S.A. Principal component analysis of 

12 growth and phenological traits explained 48.2% of the observed variability in the first principal component (PC-1). With 

multiple regressions, PC-1 was compared to environmental 

values at each source location. Regression analysis identified 

a four-variable model containing elevation, minimum January 

temperature, maximum October temperature, and February 

precipitation that explained 86% of the variability in PC-1 

(r2 = 0.86, p < 0.0001). Spatial analysis using this regression 

model identified patterns of genetic diversity within the Pacific 

Northwest that can help guide germplasm selection (i.e., seed 

collections) for restoration and revegetation activities.

Link: http://www.fs.fed.us/wwetac/threat_map/seed_zones/

Horning%20holodiscus%20genecology.pdf

667. Howe GT, Aitken SN, Neale DB, Jermstad 

KD, Wheeler NC, Chen THH. 2003. From genotype 

to phenotype: unraveling the complexities of cold adaptation in 

forest trees. Canadian Journal of Botany 81:1247-1266.

Type: Journal 

Geographic Area: North America

Keywords: association genetics, cold hardiness, dormancy, 

genecology, bud phenology, quantitative trait loci

Abstract: Adaptation to winter cold in temperate and boreal 

trees involves complex genetic, physiological, and developmental processes. Genecological studies demonstrate the 

existence of steep genetic clines for cold adaptation traits in 

relation to environmental (mostly temperature related) gradients. Population differentiation is generally stronger for 

cold adaptation traits than for other quantitative traits and 

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238 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

ability to detect evidence for adaptation depended on grazing 

regime. In addition, we measured the direct effects of grazing 

on seedling growth and survival concurrent with our investigation of local adaptation. We detected a home-site advantage 

for seedling growth or survival in both species, but the factors 

contributing to adaptive differentiation were species specific. 

Evidence of local adaptation was detected for seedling biomass in Bromus and for survivorship in Elymus. The home-site 

advantage observed in both species was greatly reduced under 

grazed conditions and in Elymus was significant only in the 

ungrazed plots. Climate and soil analyses detected significant 

differences between locations in five soil attributes and two 

climate variables. In particular, differences in exchangeable 

magnesium indicated that one of the two transplant locations 

consisted of serpentine soil, which is widely known to drive 

adaptation in plant populations. Together, these results suggest 

that it is possible to investigate the scale and factors involved in 

local adaptation with short-term transplant studies.

671. Hultine KR, Marshall JD. 2000. Altitude trends in 

conifer leaf morphology and stable carbon isotope composition.

Oecologia 123:32-40.

Type: Journal 

Geographic Area: Idaho, Montana, USA

Keywords: carbon isotope ration, stomatal density, leaf nitrogen content, leaf mass per area, evergreen conifers

Abstract: The natural ratio of stable carbon isotopes 

(δ13C) was compared to leaf structural and chemical 

characteristics in evergreen conifers in the north-central 

Rockies, United States. We sought a general model that 

would explain variation in δ13C across altitudinal gradients. Because variation in δ13C is attributed to the shifts 

between supply and demand for carbon dioxide within the 

leaf, we measured structural and chemical variables related to supply and demand. We measured stomatal density, 

which is related to CO2 supply to the chloroplasts, and leaf 

nitrogen content, which is related to CO2 demand. Leaf 

mass per area was measured as an intermediate between 

supply and demand. Models were tested on four evergreen 

conifers: Pseudotsuga menziesii, Abies lasiocarpa, Picea engelmannii, and Pinus contorta, which were sampled across 

1,800 m of altitude. We found significant variation among 

species in the rate of δ13C increase with altitude, ranging from 0.91‰ km–1 for A. lasiocarpa to 2.68‰ km–1 for 

Pinus contorta. Leaf structure and chemistry also varied 

with altitude: stomatal density decreased, leaf mass per 

area increased, but leaf nitrogen content (per unit area) 

was constant. The regressions on altitude were particularly robust in Pinus contorta. Variables were derived to 

describe the balance between supply and demand; these 

variables were stomata per gram of nitrogen and stomata 

per gram of leaf mass. Both derived variables should be 

669. Hufford KM, Mazer SJ. 2011. Local adaptation 

and the effects of grazing on the performance of Nassella pulchra: 

implications for seed sourcing in restoration. Restoration Ecology 

20:688-695.

Type: Journal 

Geographic Area: California, USA

Keywords: California grassland, home-site advantage, reciprocal transplant

Abstract: The use of local seed sources for revegetation is 

accepted practice to reduce the potential that propagules will 

be poorly adapted to site conditions. However, data are often 

lacking to determine the distance within which seed sources 

represent local genotypes. Short-term reciprocal transplant studies represent a class of tools to detect local adaptation of target 

species. We conducted a reciprocal transplant of Nassella pulchra between two central California locations to test for adaptation to local environmental conditions over a 3-year period. 

Experimental plots at one location were split between grazed 

and ungrazed sites to evaluate the potential influence of livestock grazing on the detection or magnitude of local adaptation. 

During each year of the study, evidence of a home site advantage 

depended on the location, traits studied, and population. At the 

end of the 3-year study period, however, we detected consistent 

evidence of a home-site advantage for seedling biomass among 

grazed sites at one location and ungrazed plots at the other location. In effect, local adaptation was only apparent in the final 

year of the study. Short-term reciprocal transplant studies are an 

effective tool to guide the selection of seed sources most likely 

to germinate and to become established at a restoration site, but 

such studies cannot rule out local adaptation, which may not be 

immediately detectable.

670. Hufford KM, Mazer SJ, Camara MD. 2008.

Local adaptation and effects of grazing among seedlings of two 

native California bunchgrass species: implications for restoration.

Restoration Ecology 16:59-69.

Type: Journal 

Geographic Area: California, USA

Keywords: Bromus carinatus, California grassland, Elymus 

glaucus, grazing, local adaptation, reciprocal transplant

Abstract: Adaptation to environmental factors may influence 

the germination and establishment of focal species in ecological restoration. Reciprocal transplants remain one of the best 

methods to detect local adaptation, but long-term studies are 

often not feasible. We conducted reciprocal transplants of the 

native California bunchgrasses Elymus glaucus and Bromus 

carinatus between two central California locations to seek 

evidence of adaptation to local environmental conditions in a 

single growing season. Experimental plots at one location included grazed and ungrazed sites. The combination of locations 

and grazing treatments allowed us to determine whether the 

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 239

Type: Journal 

Geographic Area: Japan

Keywords: revegetation, genetic differentiation, chloroplast 

DNA (cpDNA), amplified fragment length polymorphisms 

(AFLPs), PCR-RFLP

Abstract: We investigated the genetic structure of Miscanthus 

sinensis ssp. condensatus on Miyake Island, which was devastated by a volcanic eruption in 2000, by amplified fragment 

length polymorphisms (AFLPs) and polymerase chain reaction 

restriction fragment length polymorphism (PCR-RFLP) for 

chloroplast DNA (cpDNA) variation, to develop recommendations for the revegetation of devastated sites. Genetic differentiation among populations was significant, and five populations 

were classified into three regional groups. The aspect ratios of 

leaf-blades varied significantly among populations, but both 

geographical proximity and morphological similarities did not 

precisely reflect genetic similarities. In the airport population, 

we found a rare haplotype that may have been transmitted from 

outside the island. These findings will assist the revegetation of 

the island.

674. Iwata H, Kamijo T, Tsumura Y. 2006. Assessment 

of genetic diversity of native species in Izu Islands for a discriminate 

choice of source populations: implications for revegetation of 

volcanically devastated sites. Conservation Genetics 7:399-413.

Type: Journal 

Geographic Area: Japan

Keywords: AFLPs, Alnus sieboldiana, cpDNA, Miscanthus sinensis ssp. condensatus, Polygonum cuspidatum var. terminalis

Abstract: In using native species for revegetation, it is necessary choose source populations carefully to reduce the risk of 

planting suboptimal germplasm. To make preliminary recommendations for native species to use in the revegetation of a 

volcanically devastated area on Miyake Is., Japan, we investigated the genetic variation of Alnus sieboldiana, Miscanthus 

sinensis ssp. condensatus, and Polygonum cuspidatum var. terminalis in the Izu Islands and on the Izu Peninsula based on 

chloroplast DNA (cpDNA) sequence variations and amplified 

fragment length polymorphisms (AFLPs). The amount and 

pattern of differentiation differ between organelle and nuclear 

markers, suggesting the necessity of evaluation based on both 

types of markers. Within-population diversity did not vary 

among populations, suggesting that it does not need to be considered in the choice of a source population. The pattern and 

degree of differentiation varied among species, and geographical proximity did not necessarily accord with genetic similarity, suggesting that the site of an appropriate source population 

varies among species and should be determined empirically 

rather than by assuming that close proximity predicts genetic 

similarity. The Izu Peninsula populations deviated from the 

island populations in all species. Comparison of cpDNA sequences with those of related species indicates the possibility 

positively related to internal CO2 supply and thus negatively related to δ13C. As expected, both derived variables 

were negatively correlated with δ13C. In fact, the regression on stomatal density per gram was the best fit in the 

study (r2 = 0.72, P < 0.0001); however, the relationships 

were species specific. The only general relationship observed was between δ13C and LMA: δ13C (‰) = –32.972 

+ 0.0173×LMA (r2 = 0.45, P < 0.0001). We conclude 

that species specificity of the isotopic shift indicates that 

evergreen conifers demonstrate varying degrees of functional plasticity across environmental gradients, while the 

observed convergence of δ13C with LMA suggests that 

internal resistance may be the key to understanding interspecific isotopic variation across altitude.

672. Isik F, Keskin S, McKeand SE. 2000. Provenance 

variation and provenance-site interaction in Pinus brutia Ten.: 

consequences of defining breeding zones. Silvae Genetica 

49:213-223.

Type: Journal 

Geographic Area: Turkey, Mediterranean

Keywords: Pinus brutia, provenance variation, genotype-environment interaction, seed transfer zones, stability parameters

Abstract: Forty-nine provenances of Pinus brutia were tested 

on 13 sites in Turkey, and 10-year results were evaluated. 

Provenances did not differ for survival except at two sites. There 

were highly significant differences among the provenances in 

height and diameter in all test sites located in the Mediterranean, 

Marmara and in the southeastern regions of Turkey. However, 

provenances did not differ in Aegean region sites except for 

diameter at one site. The fastest growing provenances had up 

to 55% greater height and 50% greater diameter than the site 

means, implying considerable gain can be realized if the best 

provenances were selected for plantations. Provenances from the 

middle elevation zone (400 m to 900 m) of the Mediterranean 

region had greater growth than the provenances from peripheral 

distribution of the species at most sites. In general, the results did 

not support seed transfer zoning which was based on geographic 

and climatic data. Sub zoning of the Aegean region was not justified, as type B provenance correlations were high, well over the 

threshold value (rB = 0.8). For northern Marmara (Kesan site) 

and southeast Anatolia regions, a land race should be developed. 

Provenances were significantly different for stability variances, 

indicating the potential to select for stable genotypes across sites 

or for genotypes that are the most productive at specific sites. 

Link: http://www.silvaegenetica.de/fileadmin/content/dokument/archiv/silvaegenetica/49_2000/49-4-5-213.pdf

673. Iwata H, Kamijo T, Tsumura Y. 2004. Genetic 

structure of Miscanthus sinensis ssp. condensatus (Poaceae) on 

Miyake Island: implications for revegetation of volcanically devastated sites. Ecological Research 20:233-238.

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CHAPTER 4 • TRANSFER GUIDELINES AND ZONES • RESEARCH

climate, taking into consideration future distributions, adaptation of populations, and variability among General Circulation 

Models (GCM).

Link: http://bcwildfire.ca/ftp/HTI/external/!publish/Interim_Measures/Lw_Review_March_19_2010/INTERIM_Lw_

Report_(Jaquish)_FINAL.pdf

677. Jenkinson JL. 1975. Increasing planting stock size by 

family selection in California ponderosa pine. Berkeley (CA): 

USDA Forest Service, Pacific Southwest Forest and Range 

Experiment Station. Research Paper PSW-108. 14 p.

Type: Government Document

Geographic Area: California, USA

Compilers’ Keywords: Pinus ponderosa, transfer guidelines, 

forest management, tree improvement, genetic adaptation

Summary: This paper describes family differences in 1- and 

2-year nursery growth of ponderosa pine from natural stands 

in the northern Sierra Nevada. Wind-pollinated seed was obtained from 48 parent trees selected in eight stands. The stands 

sampled four geographic areas-three west and one east of the 

Sierra crest-to represent climates typical for ponderosa pine. In 

each area, one stand was selected on a fertile soil and the other 

on infertile ultramafic soil.

Link: http://www.fs.fed.us/psw/publications/documents/

psw_rp108/psw_rp108.pdf

678. Jenkinson JL, Nelson JA, Huddleston ME. 1993.

Improving planting stock quality—the Humboldt experience.

Albany (CA): USDA Forest Service, Pacific Southwest Research 

Station. PSW-GTR-143. 238 p.

Type: Government Document

Geographic Area: Western USA

Keywords: artificial regeneration, nursery management, plantation establishment, reforestation, seedling culture, seedling root 

growth capacity, seedling survival, Abies concolor, A. grandis, A. 

magnifica var. shastensis, A. procera, Libocedrus decurrens, Picea 

stichensis, Pseudotsuga menziesii var. menziesii, Thuja plicata, 

Tsuga heterophylla

Abstract: A seedling testing program was developed to improve the survival and growth potential of planting stock produced in the USDA Forest Service Humboldt Nursery, situated 

on the Pacific Coast in northern California. Coastal and inland 

seed sources of Douglas-fir and eight other conifers in the 

Pacific Slope forests of western Oregon and northern California 

were assessed in both nursery and field studies. Seedling top 

and root growth capacities were evaluated just after lifting and 

after cold storage, and stored seedlings were tested for survival and growth on cleared planting sites in the seed zones 

of origin. Safe lifting and cold storage schedules were defined, 

and seedling cultural regimes were formulated to produce successful 1-0, 1-1, and 2-0 stock types. Testing demonstrated the 

of hybridization with related species on the Izu Peninsula, suggesting that seeds collected from populations where related 

species live sympatrically should not be used for revegetation. 

These findings indicate the need to assess the genetic diversity 

empirically by using organelle and nuclear markers to avoid 

unintended consequences of genetic mixing associated with 

revegetation.

675. Jacobs DF, Davis AS. 2005. Genetic considerations 

in the operational production of hardwood nursery stock in the 

eastern United States. Native Plants Journal 6:4-13.

Type: Journal 

Geographic Area: Eastern USA

Keywords: tree improvement, seed zones, seed orchards, forestry, timber production, restoration

Abstract: A recent survey of forest tree nurseries in the eastern U.S. indicated that hardwood tree improvement is not 

extensively practiced at an operational level, with only 6.8% 

of hardwood seedlings produced from improved materials 

(compared to 36% for conifer seedlings at those nurseries 

that produce both). Fine hardwoods represent less than 20% 

of improved hardwood seedling production. Most respondents 

indicated that the use of genetically improved materials would 

benefit forestry in their region; however, less than 40% have 

germplasm of hardwood species in improvement programs. 

Because most respondents stated their intention to use more 

genetically improved hardwood material in seedling production over the next 10 y, availability of improved materials will 

likely limit future use. More integration of research involving 

genetic improvement into operational nursery production will 

help sustain the future value and supply of our hardwood forest 

resource.

676. Jaquish B. 2010. Review of British Columbia’s 

contemporary and projected western larch seed planning zones in 

light of climate change. Vernon, British Columbia, Canada: BC 

Ministry of Forests and Range. 1-18 p.

Type: Government Document

Geographic Area: Western Canada

Compilers’ Keywords: Larix occidentalis, bioclimatic model, 

GCM, seed zones, seed transfer, management

Background: The objectives of this range-wide project were 

to: (1) define the climate profile with a bioclimatic model that 

predicts the presence or absence of western larch from climate 

variables; (2) develop models of genetic variation that predict 

genetic differences among populations from inhabited climate; 

(3) develop and map seed zones within predicted distributions 

for present and future climates; (4) identify populations that are 

likely to become threatened and identify appropriate conservation strategies; and, (5) develop management strategies for the 

transfer of the seed sources to future location of their optimal 
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 241

CHAPTER 4 • TRANSFER GUIDELINES AND ZONES • RESEARCH

Type: Journal 

Geographic Area: Norway, Europe

Compilers’ Keywords: Norway spruce, frost hardiness

Abstract: We have compared bud set and frost hardiness 

among Norway spruce (Piceas abies (L.) Karst.) provenances 

and families in two cold-acclimation regimes in a phytotron; 

low light intensity and high night temperatures (LL–HNT), and 

high light intensity and low night temperatures (HL–NT) under 

shortening day lengths. Nine provenances from 59–66° N and 

altitude 100–700 m within Norway, and nine open-pollinated 

families from a single stand (61° N, 270 m elevation) were 

used. Both provenances and families started bud set and frost 

hardening earlier in LL–HNT than in HL–LNT. Correlations 

between the same trait expressed in two regimes were high for 

both bud set and hardiness at the provenance level and slightly 

lower at the family level. The variation among family means 

in bud set and hardiness was large. The differences found between the family extremes were up to 75% of those found between provenance extremes. The relationship between bud set 

and frost hardiness was strong among the provenance means 

within both environments (r = 0.92) but weak for the families 

(r = 0.22–0.44). Causal factors influencing phenotypic variation within traits and covariation among traits may differ for 

provenances and families within stands. The strong relationships among traits that are found at the provenance level cannot be generalized to the levels of families or clones.

681. Johnson RC, Cashman MJ, Vance-Borland K. 

2012. Genecology and seed zones for Indian ricegrass collected 

in the southwestern United States. Rangeland Ecology & 

Management 65:523-532.

Type: Journal 

Geographic Area: Southwestern USA

Keywords: Achnatherum hymenoides, common garden, genetic resources, germplasm, restoration, revegetation

Abstract: Indian ricegrass (Achnatherum hymenoides

[Roemer & J.A. Schultes] Barkworth) is a widely distributed, highly desirable native species in desert ecosystems 

in the western United States. Yet there are no studies linking genetic variation in Indian ricegrass with climate across 

major areas of its natural distribution. In this study, seeds 

from 106 collection locations from the southwestern United 

States were established in common gardens and four phenological traits (Phen; such as blooming date), six production traits (Pro; such as dry weight), and eight morphology 

traits (Morph; such as leaf dimensions) were measured in 

2007 and 2008. Analyses of variance revealed that all basic garden traits differed among source locations (P < 0.01), 

indicating widespread genetic variation. Within Phen, Pro, 

and Morph categories, canonical correlation was completed 

between basic garden traits and source location temperature 

and precipitation. This resulted in six significant (P < 0.01) 

critical elements of reforestation and proved that rapid establishment is attainable on diverse sites. Accomplishments of 

the Humboldt program recommend similar programs for other 

forest nurseries and their service regions.

Link: http://www.treesearch.fs.fed.us/pubs/24160

679. Jensen KB, Mott IW, Robins JG, Waldron BL, 

Nelson M. 2012. Genetic improvement and diversity in Snake 

River wheatgrass (Elymus wawawaiensis) (Poaceae: Triticeae).

Rangeland Ecology & Management 65:76-84.

Type: Journal 

Geographic Area: Western USA

Keywords: forage quality, genetic diversity, germplasm improvement, seedling establishment

Abstract: With the increased emphasis on using native plant 

materials in range revegetation programs in the western United 

States it is critical to identify genetically similar groups and develop native grasses that are competitive with invasive weeds, 

easy to establish, and persistent, and that produce high seed 

yield. A grass that shows appreciable drought tolerance on arid 

rangelands is Snake River wheatgrass (Elymus wawawaiensis

J. Carlson & Barkworth). This study was designed to estimate 

genetic relationships and underlying genetic components for 

seed and forage trait improvement between plant introductions (PIs) of Snake River wheatgrass, 28 half-sib Snake River 

wheatgrass families (HSFs), and cultivars Secar and Discovery 

at Nephi, Utah, between 2005 and 2006. Based on molecular genetic diversity data in Snake River wheatgrass, with the 

exception of the PIs originating from Enterprise, Oregon, all 

other collections and cultivars are not genetically different 

and represent a common gene pool from which to develop 

improved Snake River wheatgrass germplasm. Selection in 

Snake River wheatgrass for total seed yield (g/plot), 100 seed 

weight (g), and seedling emergence from a deep planting depth 

had a positive effect. Further increases through selection and 

genetic introgression from hybridization with PIs will likely 

increase seed yield and 100-seed weight, but will not increase 

seedling emergence. Increases in dry matter yield (DMY) were 

observed after two cycles of selection in the HSFs compared 

to the PIs. There remains considerable genetic and phenotypic 

variation to further increase DMY in Snake River wheatgrass 

through selection and hybridization. Trends in forage nutritional quality were not observed after two cycles of selection 

in the HSFs or the PIs and will not likely result in improvement. Through recurrent selection, populations of Snake River 

wheatgrass have been and can be developed to more effectively establish and compete on annual weed–infested rangelands.

680. Johnson O, Skroppa T. 2000. Provenances and 

families show different patterns of relationship between bud 

set and frost hardiness in Picea abies. Canadian Journal of 

Botany 30:1858-1866.
242 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

683. Johnson RC, Hellier B, Cashman M, Shaw 

NL, Erickson VJ, St. Clair JB, Vance-Borland KW. 

2010. Genecology and seed zones for native grasses and forbs. 

In: National Native Seed Conference: Native Plant Materials for 

Development, Production, and Use in Habitat Restoration. 

Corvallis (OR): Institute for Applied Ecology. 31 p.

Type: Presentation

Geographic Area: USA

Compilers’ Keywords: policy, germplasm

Abstract: Western landscapes are managed to a large extent 

for wildlife habitat, recreation, and grazing resources. However, 

urbanization, frequent fires and increasing pressure from invasive weeds are interacting to cause an unsustainable cycle 

of environmental degradation to plant communities and ecosystems. Restoration efforts that emphasize native germplasm 

adapted to “local” environments may be the best approach to 

ensure long-term sustainable management. Developing seed 

zones is a way to ensure that plant materials used for revegetation match the local environment, and are neither too broad 

nor unnecessarily local. Ongoing work with native grasses and 

forbs for conservation and seeds zone development has shown 

1) extensive genetic variation across the landscape for plant 

traits associated with growth and development, 2) growth and 

development traits are linked to climate variables at seed source 

locations, suggesting adaptive variation, 3) genetic variation in 

plant traits can be used with climate data to develop manageable seed zones to guide restoration.

Link: http://nativeseed.info/2010/presentations/JohnsonRC.pdf

684. Joly RJ, Adams WT, Stafford SG. 1989.

Phenological and morphological responses of mesic and dry site 

sources of coastal Douglas-fir to water deficit. Forest Science 

35:987-1005.

Type: Journal 

Geographic Area: Oregon, USA

Keywords: Pseudotsuga menziesii, adaptation, genetic variation, growth, water stress

Abstract: Patterns of genetic variation in morphological 

and phenological responses to water deficit are described in 

seedling progeny of four populations of coastal Douglas fir 

(Pseudotsuga menziesii [Mirb.] Franco var. menziesii), sampled 

from coastal and inland sites in Oregon. Progeny of coastal 

and inland populations differed for all traits measured, and 

differences appear to reflect adaptation to the source environment. Inland populations were characterized by early budset, 

slower rates of shoot extension, and higher root-shoot ratios. 

A significant fraction of total variability was attributable to 

differences among families within populations. Little evidence for irrigation x population interaction was found when 

seedling traits were examined singly. Of 16 traits analyzed, 

canonical variates (Phen 1, Pro 1 and 2, and Morph 1, 2, and 

3) representing each category of traits. Linear correlations (r 

> ± 25, P < 0.01) consistently linked monthly temperature 

at collection locations with Phen 1, Pro 1, and Morph 1. For 

precipitation, however, correlations were more dependent 

on month, with the strongest correlations during the spring 

developmental period. Using regression models between 

traits and climate, a map with 12 seed zones was developed 

representing much of the southwestern United States. This 

generally distinguished genetic variation between cooler and 

warmer regions, usually separating more northern, higher elevation areas from more southern, lower elevation areas. The 

correspondence between climate and genetic variation suggested climate-driven differences in natural selection, likely 

leading to adaptation. The seed zone map is recommended to 

guide and broaden germplasm collection and utilization for 

Indian ricegrass restoration.

Link: http://www.treesearch.fs.fed.us/pubs/42159

682. Johnson RC, Erickson VJ, Mandel NL, St. Clair 

JB, Vance-Borland KW. 2010. Mapping genetic variation 

and seed zones for Bromus carinatus in the Blue Mountains of 

eastern Oregon, USA. Botany 88:725-736.

Type: Journal 

Geographic Area: Oregon, USA

Keywords: genecology, Bromus carinatus, seed zones, plant 

adaptation

Abstract: Seed transfer zones ensure that germplasm selected for restoration is suitable and sustainable in diverse 

environments. In this study, seed zones were developed 

for mountain brome (Bromus carinatus Hook. & Arn.) in 

the Blue Mountains of northeastern Oregon and adjoining 

Washington. Plants from 148 Blue Mountain seed source 

locations were evaluated in common-garden studies at two 

contrasting test sites. Data on phenology, morphology, and 

production were collected over two growing seasons. Plant 

traits varied significantly and were frequently correlated 

with annual precipitation and annual maximum temperature at seed source locations (P < 0.05). Plants from warmer 

locations generally had higher dry matter production, longer leaves, wider crowns, denser foliage, and greater plant 

height than those from cooler locations. Regression models 

of environmental variables with the first two principal components (PC 1 and PC 2) explained 46% and 40% of the 

total variation, respectively. Maps of PC 1 and PC 2 generally corresponded to elevation, temperature, and precipitation gradients. The regression models developed from PC 

1 and PC 2 and environmental variables were used to map 

seed transfer zones. These maps will be useful in selecting 

mountain brome seed sources for habitat restoration in the 

Blue Mountains.

Link: http://www.treesearch.fs.fed.us/pubs/40063

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 243

Population differentiation was greatest before the first frost 

however, relative hardiness of populations before and after 

the first frost were strongly correlated. Hardiness was related 

to latitude, longitude, and elevation of population origins. 

Multiple regression analyses accounted for 74% of the variance among population means. Population differentiation was 

detected across relatively small geographic and elevational intervals. Consequently, in order to limit maladaptation, transfer 

of plant material in artificial reforestation should be carefully 

controlled. 

687. Joyce D, Rehfeldt GE. 2013. Climatic niche, 

ecological genetics, and impact of climate change on eastern white 

pine (Pinus strobus L.): guidelines for land managers. Forest 

Ecology and Management 295:173-192.

Type: Journal 

Geographic Area: Ontario, Canada

Keywords: bioclimate models, ecological genetics, climate 

niche, climate response models, seed transfer guidelines, seed 

zones

Abstract: Approximately, 123,500 forest inventory and ecological ground plots representing eastern North America were 

used to predict the contemporary distribution of eastern white 

pine (Pinus strobus L.) from climate. The random forests classification tree procedures produced an 8-variable algorithm 

that had an 8% overall error rate. Erroneous predictions of 

presence, or errors of commission, were 13%, while falsely 

predicting absence, or errors of omission were 1%. Climatebased multiple regression models were developed to describe 

patterns of genetic variation among 112 populations representing the range of P. strobus in Ontario, Canada east of Lake 

Superior. Degree days > 5 °C was the best predictor of variation in phenology and growth potential, with 5-year height providing the greatest resolution of inter-population variation (R2

= 0.68). Cold hardiness in the fall was most closely associated 

with mean minimum temperature (R2 = 0.26). Height growth 

data from four disparate provenance test series that together 

included a total of 354 provenances corroborated the rangewide applicability of the regional genetic models. Although 

variation in growth potential in the central Appalachian 

Mountains was most closely associated with mean minimum 

temperature, degree days > 5 °C remained the best predictor of 

range-wide variation in growth potential (R2 = 0.41). The contemporary distribution and inter-population genetic variation 

were projected into future climates predicted by three General 

Circulation Models, two scenarios, and three time steps. All 

projections indicate early and sustained deterioration in the 

contemporary habitat. Concurrence among projections regarding the redistribution of suitable habitat to the north of the contemporary distribution identifies geographic locations with the 

highest probability of supporting vigorous stands of P. strobus. 

Concurrences among genetic projections clarify the intraspecific redistribution required to conserve adaptive variation. The 

only average daily rate of growth and seedling height had significant interaction terms. Interactions between irrigation and 

families-within-population were more prevalent. A canonical 

discriminant analysis was used to identify a subset of variables 

that best reveals differences among progeny of coastal and inland populations. Implications for seed transfer and for selection and breeding of genotypes suitable for xeric environments 

are discussed.

685. Jonsson A, Eriksson G, Franzen A. 1986.Withinpopulation variation in frost damage in Pinus contorta Dougl. 

seedlings after simulated autumn or late-winter conditions. Silvae 

Genetica 35:96-102.

Type: Journal 

Geographic Area: Canada, Sweden

Keywords: Pinus contorta, climate chamber, frost damange, 

within-population variation

Summary: Single-tree progenies from a few populations of 

Pinus contorta were cultivated in a climate chamber and tested 

with respect to frost tolerance. The plants were exposed to 

-10 °C for three hours during the hardening period, simulating autumn conditions. Single-tree progenies and population 

samples were exposed to two, six, or twelve large diurnal temperature fluctuations, -10 to +20 °C, simulating conditions inducing dehardening during late winter. Significant differences 

in frost damage were found between populations as well as between single-tree progenies from one of the populations after 

freezing during autumn conditions. The family repeat abilities 

for frost damage exceeded 0.50 in two of the three populations 

studied. Damage to plants increased with increasing number 

of simulated late-winter temperature fluctuations. This was the 

case both for the roots and the upper parts of the plants. No significant within-population variation in frost damage following 

exposure to simulated late-winter conditions as obtained. The 

simulating technique used is laborious and will not be further 

developed. There was a non-significant, positive relationship 

between frost damage induced in single-tree progenies by 

simulated autumn and late-winter conditions. 

Link: http://www.silvaegenetica.com/fileadmin/content/dokument/archiv/silvaegenetica/35_1986/35-2-3-96.pdf

686. Joyce DG. 1988. Adaptive variation in cold hardiness of eastern larch, Larix laricina, in northern Ontario.

Canadian Journal of Botany 18:85-89.

Type: Journal 

Geographic Area: Ontario, Canada

Compilers’ Keywords: local adaptation, cold hardiness, transfer guidelines

Abstract: Adaptive differentiation in cold hardiness of Larix 

laricina (DuRoi) K. Koch from 66 populations, primarily from 

northern Ontario, was studied using laboratory freezing tests. 

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Therefore, insights into intraspecific variation within and 

among tree populations in climate response are of paramount 

importance. In our study we investigated intraspecific variation in climate response among Norway spruce (Picea abies) 

populations in the eastern Alpine range. Results from a comprehensive Austrian provenance test, comprising tree heights 

at age 15 from 379 populations planted at 29 test sites across 

Austria, were used to calibrate climate response functions for 

groups of Norway spruce populations. Potential future changes 

in productivity for climate change conditions as represented 

by a regionalized A1B scenario were estimated using height 

at age 15 as a productivity proxy. Climate response functions 

were calculated for single populations and aggregated clusters 

of populations from climatically similar origins. Our results 

hardly revealed any declines in employed proxies for productivity of Norway spruce throughout its current distribution 

range in Austria. For most parts of Austria an increase of tree 

heights up to 45 percent can be expected until 2080. However, 

the impact of a warming climate is different for individual 

population groups. Generally, variation in climate response increases with higher temperatures and less precipitation. Thus, 

an optimized choice of seed material according to prospective 

future climate conditions has the potential for an additional increase of productivity up to 11 percent. In general, populations 

from currently warm and drought prone areas seem to be well 

adapted to respective climate conditions and may be appropriate candidates for extended utilization in future. Furthermore, 

populations showing the best productivity indices originate 

from regions, which are phylogenetically distinct from the core 

distribution area of Norway spruce, suggesting that population 

history might explain part of the variation in climate response 

among populations.

Link: http://www.researchgate.net/publication/220039116_

Intraspecific_variation_in_climate_response_of_Norway_spruce_in_the_eastern_Alpine_range_Selecting_appropriate_provenances_for_future_climate/file/

d912f4fdeea8cb6ea9.pdf

690. Keir KR, Bemmels JB, Aitken SN. 2011. Low 

genetic diversity, moderate local adaptation, and phylogeographic 

insights in Cornus nuttallii (Cornaceae). American Journal of 

Botany 98:1327-1336.

Type: Journal 

Geographic Area: USA

Keywords: adaptive cline, chloroplast diversity, Cornaceae, 

Cornus nuttallii, microsatellite markers, Pacific dogwood, phenotypic variation, population structure, postglacial recolonization

Abstract: Genetic knowledge is completely lacking for 

Pacific dogwood (Cornus nuttallii), a western North American 

temperate tree that is pollinated and dispersed by biological 

vectors. We investigated how history, geography, and climate 

have affected population genetic structure, local adaptation, 

projections have direct relevance in developing management 

strategies for accommodating the changing climate.

Link: http://forest.moscowfsl.wsu.edu/climate/JoyceRehfeldt2013.pdf 

688. Kabrick JM, Dey DC, Gwaze D. 2007. Shortleaf 

pine restoration and ecology in the Ozarks: proceedings of a 

symposium. Delaware (OH): USDA Forest Service, Northern 

Research Station. General Technical Report NRS-P-15. 224 p.

Type: Government Document

Geographic Area: Southeastern USA

Compilers’ Keywords: Pinus echinata, transfer zones, seed 

zones

Foreward: The Shortleaf Pine Restoration and Ecology in the 

Ozarks Symposium was held at the University Plaza Hotel 

and Convention Center in Springfield, Missouri, on November 

7-9, 2006. The purpose of the symposium was to communicate experiences, research, successes, challenges, and inspire 

inquiries into the ecology, management, and restoration of 

shortleaf pine communities and ecosystems in which shortleaf 

pine is prominent. The symposium brought together more than 

200 registrants including private landowners, state and federal resource managers, consultants, members of conservation 

groups, and scientists from public and private institutions to 

exchange ideas and share knowledge about ecosystems containing shortleaf pine. Altogether, the symposium featured 

eight plenary presentations, 36 oral presentations, and nine 

poster presentations.

Link: http://www.treesearch.fs.fed.us/pubs/12785

689. Kapeller S, Lexer MJ, Geburek T, Hiebl J, 

Schueler S. 2012. Intraspecific variation in climate response 

of Norway spruce in the eastern alpine range: selecting appropriate 

provenances for future climate. Forest Ecology and Management 

271:46-57.

Type: Journal 

Geographic Area: Austria, Europe

Keywords: climate change impacts, Gaussian response model, 

quantitative genetic variation, provenance test, Picea abies

Abstract: Enhancing adaptation of forest ecosystems to prospective climate change is a major challenge in current forest management. Beyond potential negative effects of climate 

change such as decreasing productivity due to an increasing 

number of drought periods and damages from intensified 

disturbance regimes, there is also a potential for increasing 

productivity due to prolonged vegetation periods and higher 

photosynthetic rates. Quantitative genetic variation is crucial 

for adaptability of species towards environmental changes. 

The use of suitable reproductive material for forest regeneration will be a key factor essential for both, mitigating negative effects and making the most of potential positive effects. 

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 245

with site, using southern sources. Results at PS strongly support the use of local seed, with allowance for transferring seed 

to adjacent zones.

692. Kitzmiller JH. 2009. Regional genetic variation in 

three native grasses in northern California. Native Plants Journal 

10:263-280.

Type: Journal 

Geographic Area: California, USA

Keywords: geographic variation, seed transfer zones, adaptive 

traits, Elymus glaucus, Bromus carinatus, Bromus orcuttianus

Abstract: Germination, growth, and flowering traits varied 

widely in a common-garden nursery environment and expressed coarse-textured geographic patterns among Plumas 

National Forest seed sources of 3 native grasses (Poaceae): 

blue wildrye (Elymus glaucus Buckley ssp. glaucus, California 

brome (Bromus carinatus Hook. & Arn. var. carinatus), and 

Orcutt’s brome (B. orcuttianus Vasey). Seed zones are justified 

to partition the geographic variation in grasses on the Plumas 

National Forest because: 1) adaptive traits varied significantly 

with geographic surrogates for climate and photoperiod; 2) a 

coarse-textured geographic and topographic pattern overshadowed microsite variation; and 3) paired (proximal) sources 

were more similar to one another than more distal sources, all 

of which indicated selection may have contributed to population differentiation. Each species displayed a different geographic pattern, but Bromus species were most similar. Lowelevation southwestern sources of B. carinatus had earlier and 

higher germination and slightly higher shoot dry weight, while 

high-elevation northern lots had later and lower germination 

and dry weight. Southern sources of B. orcuttianus produced 

more flowers and germinated earlier than northern sources. 

Elymus glaucus from lower-elevation and more eastern sources, where summer drought is most pronounced, had higher dry 

weight and flowers.

693. Kitzmiller JH, Hanson L. 2011. Patterns of adaptation in three native grasses in northern California. Native Plants 

Journal 12:45-61.

Type: Journal 

Geographic Area: California, USA

Keywords: seed transfer zones, local adaptation, Elymus 

glaucus, Bromus carinatus, Bromus orcuttianus, Sierra Nevada, 

Poaceae

Abstract: Provisional seed transfer zones were developed 

for 3 Poaceae grasses, Elymus glaucus Buckley ssp. glaucus, 

Bromus carinatus Hook. & Arn., and Bromus orcuttianus

Vasey, from a 4-y study with 11 reciprocal-transplant gardens 

across the Plumas National Forest. To conserve existing adaptive patterns, 4 seed zones were proposed for B. carinatus and 

B. orcuttianus, and six for E. glaucus. Adaptive traits of source 

and the phylogeography of this species. We examined patterns and levels of diversity in nuclear microsatellites (SSRs) 

and cpDNA haplotypes in populations from across the species 

range. We compared these results to population differentiation 

and genetic clines in phenotypic traits in a common garden. 

Genetic diversity was low for both nuclear SSRs and cpDNA. 

There was a lack of population structure (FST = 0.090) in the 

coastal portion of the species range, with estimates of population genetic diversity in microsatellite markers decreasing 

with latitude from California to British Columbia. A disjunct 

interior population in Idaho 450 km from the coastal range had 

the lowest diversity but the highest divergence of all populations studied. Only a single nucleotide polymorphism was 

discovered after sequencing 5547 base pairs in seven noncoding regions of cpDNA. Both cpDNA haplotypes were widely 

distributed throughout the species range. Quantitative variation among populations was moderate (0.11 ≤ QST ≤ 0.63), 

and weak but significant adaptive clines were found between 

quantitative traits and population climatic variables (0.09 ≤ R2

≤ 0.34). Cornus nuttallii likely faced a population bottleneck in 

a single southern refugium during the Last Glacial Maximum. 

Despite low genetic diversity, it is weakly to moderately locally adapted.

Link: http://www.ncbi.nlm.nih.gov/pubmed/21821593

691. Kitzmiller JH. 2005. Provenance trials of ponderosa 

pine in northern California. Forest Science 51:595-607.

Type: Journal 

Geographic Area: California, USA

Keywords: tree improvement, genetics, seed zones, seed 

transfer

Abstract: Growth of 17 ponderosa pine populations transferred within the southern Cascades–Sierra Nevada ranges 

were tested for 19 years on two contrasting sites, Cherry Hill 

(CH) and Peanut Springs (PS). Seven additional southern 

Sierra populations were tested at PS. Results provide guidance to forest managers seeking to reforest ponderosa pine 

after large wildfires or to improve productivity of plantations. PS represented warm, moist, lower mixed-conifer sites 

in the central Sierra. CH represented colder, drier climates at 

the southern limit of the Cascades in upper elevation mixed 

conifer. High inherent site productivity at PS was reduced by 

topsoil displacement and brush competition for moisture and/

or nutrients. Contrasting magnitudes and patterns of growth 

among provenances were expressed between sites. The local 

provenance at PS outgrew more distant sources by 12 to 74%. 

However, high elevation, nonlocal Sierra sources grew best 

at CH, where volume decreased with source latitude and increased with source elevation. All sources had stockier form at 

CH. In both tests, Sierra sources outgrew and were stockier than 

Cascade sources; stockiness increased with source elevation. 

Current results at CH support matching provenance elevation 

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246 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

glaucus may need to be collected in close proximity to the proposed restoration site.

695. Knapp EE, Rice KJ. 1997. Ecotypes of native species: 

how local is local in restoration plantings? In: Kelly M, Wagner 

E, Warner P, editors. California Exotic Pest Plant Council 1997 

Symposium Proceedings Volume 3. Trabuco Canyon (CA): 

California Exotic Pest Plant Council. p 1-5.

Type: Conference Paper

Geographic Area: California, USA

Compilers’ Keywords: local adaptation, native grasses, 

genetic diversity, Elymus glaucus, Nassella pulchra, reciprocal transplant

Conclusion: Populations of native species growing in different environments and separate regions are often genetically 

distinct. These genetic differences should be considered when 

source material of native species is obtained for restoration 

and revegetation. We investigated patterns of genetic variation 

for different types of traits among populations of a popularly 

planted native grass (Nassella pulchra), and determined the extent to which local populations of Elymus glaucus, and N. pulchra are better adapted than non-local populations. We found 

strong genetic differences among populations of N. pulchra for 

both isozyme markers and quantitative traits, but the patterns 

of genetic differentiation visualized by the two methods were 

not similar. Thus, management recommendations and seed 

transfer guidelines should not be based on data for only one 

type of trait. In addition, patterns of quantitative trait variation 

in N. pulchra were strongly correlated with climatic variation 

among sites, suggesting that it may be possible to obtain a 

rough match between seed sources and planting sites by using 

readily available climate zone data.

Link: http://www.cal-ipc.org/symposia/archive/pdf/1997_

symposium_proceedings1941.pdf

696. Knapp EE, Rice KJ. 2011. Effects of competition and 

temporal variation on the evolutionary potential of two native 

bunchgrass species. Restoration Ecology 19:407-417.

Type: Journal 

Geographic Area: California, USA

Keywords: Elymus glaucus, genetic variation, Nassella pulchra, 

phenotypic plasticity, selection response

Abstract: The capacity of restored plant populations to adapt 

to new environmental challenges depends on within population genetic variation. We examined how much genetic and 

environmentally based variation for fitness associated traits 

exists within populations of two native grasses commonly 

used for restoration in California. We were also interested in 

understanding how phenotypic expression of genetic variation 

for these traits varies with growth environment. Thirty maternal families of Elymus glaucus (Blue wild rye) and Nassella 

populations were correlated with their geographic, climatic, 

and ecologic origins. Findings based on seed source x environment interactions and the geographic-climatic patterns for 

local, distant, and proximal paired populations suggest that 

natural selective pressures have produced weak to moderate 

broad-scale local adaptation. Three consistent “coarse-textured” adaptive patterns emerged: 1) sources from mesic westside and east-side ecological zones formed two well-differentiated groups; 2) sources from the broad intermediate area (west 

side central and transition) were less differentiated and often 

intergraded with the mesic west-side (southwestern) and eastside (northeastern) groups; and 3) mesic west-side was divided 

into 2 elevation bands. Local adaptation was found less often 

on a finely tuned local scale.

694. Knapp EE, Rice KJ. 1996. Genetic structure and gene 

flow in Elymus glaucus (blue wildrye): implications for native 

grassland restoration. Restoration Ecology 4:1-10.

Type: Journal 

Geographic Area: Western USA

Compilers’ Keywords: seed zones, genetic variation, allozyme variation, electrophoresis, revegetation, native grass, 

population genetic structure, seed collection, self-pollination

Abstract: Interest in using native grass species for restoration is increasing, yet little is known about the ecology and 

genetics of native grass populations or the spatial scales over 

which seed can be transferred and successfully grown. The 

purpose of this study was to investigate the genetic structure 

within and among populations of Elymus glaucus in order to 

make some preliminary recommendations for the transfer and 

use of this species in revegetation and restoration projects. 

Twenty populations from California, Oregon, and Washington 

were analyzed for allozyme genotype at 20 loci, and patterns 

of variation within and among populations were determined. 

Allozyme variation at the species level was high, with 80% 

of the loci polymorphic and an average expected heterozygosity (an index of genetic diversity) of 0.194. All but two of the 

populations showed some level of polymorphism. A high degree of population differentiation was found, with 54.9% of 

the variation at allozyme loci partitioned among populations 

(Fst = 0.549). A lesser degree of genetic differentiation among 

closely spaced subpopulations within one of the populations 

was also demonstrated (Fst = 0.124). Self-pollination and the 

patchy natural distribution of the species both likely contribute 

to the low level of gene flow (Nm = 0.205) that was estimated. 

Zones developed for the transfer of seed of commercial conifer 

species may be inappropriate for transfer of E. glaucus germplasm because conifer species are characterized by high levels 

of gene flow. Limited gene flow in E. glaucus can facilitate the 

divergence of populations over relatively small spatial scales. 

This genetic differentiation can be due to random genetic drift, 

localized selective pressures, or both. In order to minimize 

the chances of planting poorly adapted germplasm, seed of E. 

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 247

phenotype. Hence seed quality may be associated with the paternal plant response to the environment.

Link: http://www.ncbi.nlm.nih.gov/pubmed/21434931

698. Krakowski J, Stoehr MU. 2009. Coastal Douglasfir provenance variation: patterns and predictions for 

British Columbia seed transfer. Annals of Forest Science 

66:811-821.

Type: Journal 

Geographic Area: Western Canada

Keywords: adaptation, genecology, provenance, Pseudotsuga 

menziesii, seed transfer

Abstract: We evaluated performance variability in two series 

of provenance trials of Pseudotsuga menziesii var. menziesii. EP 

480 included 16 reciprocal provenances planted from British 

Columbia to Oregon, measured up to age 45. EP 599.03 featured five common provenances, plus the local source, planted 

at 23 British Columbia sites, measured up to age 33. Although 

residual variance was high, site accounted for 31 to 60% of 

the volume variance, while provenance accounted for 1–6%. 

Interactions were also significant across ages and trials. 

Genotype environment interaction was evident in EP 599.03, 

but not EP 480, which may reflect differences in experimental 

design. The worst provenances always ranked low across sites 

and over time. No geographic or climatic (annual, seasonal, 

monthly) variables consistently explained patterns of volume 

across sites or ages, singly or in combination for either trial, 

similar to findings from other studies of coastal Douglas-fir. 

Provenances from Washington to central Oregon often performed as well as the local provenance or better at British 

Columbia trial sites. Populations from higher elevations, poor 

sites and submaritime provenances were less vigorous. Results 

support maintaining elevational and ecotypic transfer limits, 

emphasizing site-specific decision making, and permitting 

wider latitudinal transfer on similar quality sites.

699. Krauss SL, He TH. 2006. Rapid genetic identification 

of local provenance seed collection zones for ecological restoration 

and biodiversity conservation. Journal for Nature Conservation 

14:190-199.

Type: Journal 

Geographic Area: Australia

Keywords: Acacia, AFLP, genetic diversity, habitat restoration, 

home-site advantage, provenance

Abstract: The ecological restoration of native plant communities requires the collection of large amounts of seed. 

Use of non-local provenance seed can have detrimental consequences for the success of restoration if there is a home-site 

advantage, and for nature conservation through the erosion 

of natural patterns of population genetic structuring and/or 

genetic swamping (and extirpation) of locally significant 

pulchra (Purple needlegrass) were sampled from both coastal 

and interior populations and reciprocally transplanted into 

three replicated common gardens with and without interspecific competition at each site. Reproductive output of families 

differed both among years and with competition treatments. 

Phenotypic expression of genetic variation in culm production 

differed among populations and was very low when families 

were grown with interspecific competition. Without interspecific competition, the degree of genetic determination peaked 

in year two in both species (8.4 and 15.1% in E. glaucus and 

N. pulchra, respectively). Significant genetic differences in 

reproduction and phenotypic plasticity were found among N. 

pulchra subpopulations sampled less than 3 km apart, further 

highlighting the importance of thoroughly sampling available 

genetic variation in populations used for restoration. The variable and generally low expression of genetic variation indicates that rates of adaptation in restored populations of these 

native grasses may vary temporally and may be especially 

slow within competitive environments.

Link: http://www.fs.fed.us/psw/publications/knapp/

psw_2009_knapp005.pdf

697. Kochanek J, Steadman KJ, Probert RJ, Adkins 

SW. 2011. Parental effects modulate seed longevity: exploring 

parental and offspring phenotypes to elucidate pre-zygotic 

environmental influences. New Phytologist 191:223-233.

Type: Journal 

Geographic Area: Australia

Keywords: maternal, paternal environment, paternal, Plantago 

cunninghamii, pre-zygotic, seed ageing, seed longevity, 

transgenerational

Abstract: Seed longevity, which is essential for germplasm 

conservation and survival of many land plant species, can vary 

considerably within species and cultivars. Here, we explore 

the relationship between parental and offspring phenotypes to 

elucidate how pre-zygotic environment affects seed longevity. 

Plants of the wild species Plantago cunninghamii were exposed to wet or dry soil within a warm or cool glasshouse until 

flowering and then moved to a common environment. Seeds 

subsequently produced were collected at maturity, and longevity was assessed by controlled ageing at 45 °C, 60% relative 

humidity. Multivariate analysis was used to examine relationships between the parental and offspring phenotypes. The 

pre-zygotic environment resulted in a highly plastic parental 

response which was passed on to offspring seeds and changed 

their longevity (p50) by more than a factor of 2. Seed longevity 

is a function of the seed population’s distribution of deaths in 

time (σ) and quality (Ki); σ was associated with plant size, 

and Ki with reproductive plant traits. The pre-zygotic growth 

environment modulated seed longevity via a parental effect. 

Reproductive performance and seed quality (Ki) were highly 

correlated with each other and unrelated to the maternal plant 

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248 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Keywords: climate change response functions, genotype by 

environment interaction, provenance tests, provenance transfer 

functions, Pseudotsuga menziesii var. glauca

Abstract: Projected climate change will affect existing forests, as substantial changes are predicted to occur during their 

life spans. Species that have ample intraspecific genetic differentiation, such as Douglas-fir (Pseudotsuga menziesii (Mirb.) 

Franco), are expected to display population-specific growth 

responses to climate change. Using a mixed-effects modeling 

approach, we describe three-year height (HT) growth response 

to changes in climate of interior Douglas-fir populations. We 

incorporate climate information at the population level, yielding a model that is specific to both species and population. 

We use data from provenance tests from previous studies that 

comprised 236 populations from Idaho, Montana, and eastern 

Washington, USA. The most sensitive indicator of climate 

was the mean temperature of the coldest month. Population 

maximum HT and HT growth response to changes in climate 

were dependent on seed source climate. All populations had 

optimum HT growth when transferred to climates with warmer 

winters; those originating in sites with the warmest winters 

were taller across sites and had highest HT growth at transfer distances closest to zero; those from colder climates were 

shortest and had optimum HT growth when transferred the 

farthest. Although this differential response damped the height 

growth differences among populations, cold-climate populations still achieved their maximum growth at lower temperatures than warm climate populations. The results highlight 

the relevance of understanding climate change impacts at the 

population level, particularly in a species with ample genetic 

variation among populations.

Link: http://www.treesearch.fs.fed.us/pubs/40383

702. Lesica P, Allendorf FW. 2002. Ecological 

genetics and the restoration of plant communities: mix or match?

Restoration Ecology 7:42-50.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: conceptual framework, native plant 

materials, hybrids, mixed species, local adaptation, cultivar

Abstract: We present a conceptual framework for choosing 

native plant material to be used in restoration projects on the 

basis of ecological genetics. We evaluate both the likelihood of 

rapid establishment of plants and the probability of long-term 

persistence of restored or later successional communities. In 

addition, we consider the possible harmful effects of restoration projects on nearby ecosystems and their native resident 

populations. Two attributes of the site to be restored play an 

important role in determining which genetic source will be 

most appropriate: 1) degree of disturbance and 2) size of the 

disturbance. Local plants or plants from environments that 

“match” the habitat to be restored are best suited to restore 

genotypes. As part of an ongoing project to genetically delineate local provenance seed collection zones for species 

within a large urban bushland remnant of high conservation 

value, we assessed population genetic differentiation in two 

widespread coastal leguminous species, Acacia rostellifera

and A. cochlearis (Fabaceae), commonly used in restoration 

programmes in SW Australia. Using amplified fragment 

length polymorphism (AFLP), we found very high levels of 

genetic differentiation among populations, with an analysis 

of molecular variance (AMOVA) showing more than 50% 

of the total genetic variance to be partitioned among populations (φst = 0.58 and 0.51 for A. rostellifera and A. cochlearis, 

respectively), and marked non-overlap of almost all potential seed source populations from the local population in 

ordinations. Our results suggest extremely restricted natural 

dispersal among populations, possibly due to a combination 

of low seed set, seed dispersal by ants, clonality, a linear distribution of naturally fragmented populations and possibly 

low outcrossing rates. We suggest a narrow seed collection 

zone should be applied to these species for the conservation of genetic diversity and natural patterns of population 

genetic structure, and we highlight the value and importance 

of provenance evaluation to ecological restoration.

Link: http://peeg-curtin.com/wp-content/uploads/2013/05/26.pdf

700. Kuser JE, Ching KK. 1980. Provenance variation 

in phenology and cold hardiness of western hemlock seedlings. 

Forest Science 26:463-470.

Type: Journal 

Geographic Area: Western North America

Keywords: Tsuga heterophylla, bud burst, bud set

Abstract: Seedlings from a range-wide sample of 20 western hemlock [Tsuga heterophylla (Raf.) Sarg.] provenances 

showed much variation in phenology and cold hardiness at 

Corvallis, Oregon. Seedlings of northern, high-elevation, or 

Rocky Mountain provenances set and burst bud earlier and 

survived early frost and winter cold better than seedlings of 

southern, low elevation, or coastal provenances. A strong 

north-south cline was found in the variation among coastal 

provenances. A sample of three families per provenance revealed four cases of marked intraprovenance variation, two 

in bud-set date and two in cold survival.

701. Leites LP, Robinson AP, Rehfeldt GE, Marshall 

JD, Crookston NL. 2012.Height-growth response to climate 

changes differs among populations of Douglas-fir: a novel analysis 

of historic data. Ecological Applications 22:154-165.

Type: Journal 

Geographic Area: Northwestern USA

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 249

Compilers’ Keywords: seed transfer guidelines, provenance, 

breeding zones

Abstract: Known patterns of genetic variation among provenances are useful in gene resource management such as guiding seed transfer and delineating breeding zones. The objectives of this study were (1) to determine the genetic structure 

of white spruce (Picea glauca (Moench) Voss) from Quebec 

and southeastern Ontario, (2) to develop models describing 

patterns of genetic variation among provenances and their relationships with environmental variation, (3) to test the validity 

of the models by using independent data sets, and (4) to guide 

seed transfer and to delineate provisional breeding zones for 

white spruce in Quebec. A sample of 63 provenances was used 

in this study, with each provenance composed of one to five 

separate families (a total of 263 families). Data were collected 

on seedling heights at various ages, 1-year branch number, 

and 3-year bud burst and bud set. Significant differences were 

found for each trait among provenances and among families 

within provenances. Provenances and families within provenances accounted for similar amounts of the genetic variability. All traits were moderately to strongly intercorrelated at the 

provenance level. Two principal components composed 87% 

of the total variation for all traits. Regression models explained 

between 19% and 65% (an average of 47%) of provenance 

variation and showed that patterns of provenance variation followed mainly a south-north cline and to a lesser extent a westeast cline. The regression models were tested using data from 

seven field tests, which showed that the models performed 

fairly well in predicting relative risk of seed transfer. White 

spruce provenances could be transferred long distances with 

small relative risk. We delineated two provisional breeding 

zones for white spruce in Quebec from the models describing 

patterns of genetic variation among provenances.

705. Li H, Wang X, Hamann A. 2010. Genetic 

adaptation of aspen (Populus tremuloides) populations to spring 

risk environments: a novel remote sensing approach. Canadian 

Journal of Botany 40:2082-2090.

Type: Journal 

Geographic Area: North America

Compilers’ Keywords: genetic variation, climate change, 

seed transfer, spring drought

Abstract: This study investigates geographic patterns of genetic variation in aspen (Populus tremuloides) spring phenology with the aim of understanding adaptation of populations 

to climatic risk environments and the practical application 

of guiding seed transfer. We use a classical common garden 

experiment to reveal genetic differences among populations 

from western Canada and Minnesota, and we present a novel 

method to seamlessly map heat-sum requirements in northern 

and high-elevation aspen populations from the central boreal 

plains of Saskatchewan and Alberta, and populations from 

sites where degree of disturbance has been low. Hybrids or 

“mixtures” of genotypes from different sources may provide 

the best strategy for restoring highly disturbed site to which 

local plants are not adapted. Cultivars that have been modified 

by intentional or inadvertent selection have serious drawbacks. 

Nevertheless, cultivars may be appropriate when the goal is 

rapid recovery of small sites that are highly disturbed.

Link: http://www.esalq.usp.br/lcb/lerf/divulgacao/recomendados/artigos/lesica1999.pdf

703. Lesser MR, Parker WH. 2006. Comparison of 

canonical correlation and regression based focal point seed 

zones of white spruce. Canadian Journal of Forest Research 

36:1572-1586.

Type: Journal 

Geographic Area: Canada

Compilers’ Keywords: Picea glauca, CANCOR, seed transfer, 

methods

Abstract: The focal point seed zone methodology determines 

spatially explicit areas of adaptive similarity for any selected 

geographic point and is used to match seed sources and planting sites. A total of 127 seed sources (provenances) of white 

spruce (Picea glauca (Moench) Voss) from Ontario and western 

Quebec were established at a greenhouse and in six field trials 

throughout Ontario. Growth and phenological variables were 

measured over three growing seasons. Two focal point seed 

zone methodologies were employed: (i) using models derived 

from principal components analysis (PCA) of biological response variables followed by multiple linear regression against 

climate variables and (ii) using models derived from canonical 

correlation analysis (CANCOR). While both approaches use 

climate data to model adaptive variation, CANCOR reduces 

the number of steps in the analysis by simultaneously finding the relationships of biological and climatic variables that 

maximize the covariance between two data sets. Although 

more of the variation in adaptive biological traits was actually described by climate variables using the PCA-regression 

approach, this method produced intuitively less realistic patterns. Both methods showed similar overall geographic trends, 

but the CANCOR method had a finer resolution, especially 

in southern Ontario, presumably due to statistical efficiency; 

growth was modeled by all climate variables. 

Link: http://flash.lakeheadu.ca/~whparker/CJFR2006.pdf

704. Li P, Beaulieu J, Bousquet J. 1997. Genetic structure and patterns of genetic variation among populations in eastern white spruce (Picea glauca). Canadian Journal of Botany 

27:189-198.

Type: Journal 

Geographic Area: Canada

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250 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Abstract: A gap analysis was conducted to evaluate the extent 

to which genetic resources are conserved in situ in protected 

areas for six conifer species in the Pacific Northwest (Sitka 

spruce, sugar pine, western white pine, ponderosa pine, western 

redcedar, and western hemlock). The gap analysis involved producing a GIS database detailing the location of protected areas 

and the distribution and abundance of tree species as inferred 

from data on potential plant association groups, actual plant associations, and actual land cover type. We used two schemes for 

stratifying the distribution of each species into genetic populations for gap analysis: seed zones and ecoregions. The results 

show that most seed zones and ecoregions contain at least 5,000 

mature individuals in protected areas, indicating strong in situ

gene conservation. Protection is less complete, however, for 

western white pine in the Puget lowlands, where urbanization 

on disease have reduced populations below safe levels. These 

populations represent the highest priority for additional gene 

conservation. Other species and areas warranting further evaluation include Sitka spruce in some parts of the Puget lowlands, 

remnant western white pine stands in the Oregon Coast Range, 

and sugar pine within the white pine blister rust zone.

Link: http://www.treesearch.fs.fed.us/pubs/29710

708. Lu P, Joyce DG, Sinclair RW. 2003. Geographic 

variation in cold hardiness among eastern white pine (Pinus strobus

L.) provenances in Ontario. Forest Ecology and Management 

178:329-340.

Type: Journal 

Geographic Area: Ontario, Canada

Keywords: Pinus strobus, cold hardiness, geographic variation, 

seed transfer

Abstract: Needle and shoot samples of eastern white pine 

provenances in Ontario were subjected to a series of artificial 

freezing tests to determine provenance differentiation in fall and 

winter cold hardiness. Results indicated that under the natural 

processes of cold acclimation in the fall, above ground tissues 

of eastern white pine could withstand low freezing temperatures 

in September–October that exceeded the local long-term minimum climate extremes of the same periods. The severity of cold 

damage increased at a faster rate in September than in October 

and November when temperature was further decreased below 

the thresholds that started to cause cold damage. Under the selected freezing temperatures that inflicted between 15 and 85% 

average needle or/and cambium damage, consistent results 

were observed among freezing tests that were conducted using 

samples of different ages and cold acclimation stages. A clinal 

pattern of geographic variation in the severity of cold damage 

was detected with a trend parallel to that of growth potential. As 

thermal conditions improve from the north to south in Ontario, 

the growth potential of a provenance increases but the degree 

of cold hardiness decreases. Based on risk assessments using 

a spatial regression approach, it was, however, believed that 

Minnesota exhibit moderately low heat-sum requirements 

for budbreak. Analysis of corresponding climate normal data 

shows that late budbreak due to both frost and drought risks in 

early spring. We therefore caution against distance seed transfer of Minnesota provenances to the boreal plains of Alberta 

and Saskatchewan. Although such transfers have been shown 

to increase tree growth in short-term field tests, this planting 

material may be susceptible to exceptional spring droughts.

Link: http://www.ualberta.ca/~xianli/pdfs/Li%20et%20

al%202010%20-%20CJFR.pdf

706. Li P, Beaulieu J, Daoust G, Plourde A. 1997. 

Patterns of adaptive genetic variation in eastern white pine (Pinus 

strobus) from Quebec. Canadian Journal of Botany 27:199-206.

Type: Journal 

Geographic Area: Canada

Compilers’ Keywords: provenance, progeny test, genetic 

variation, genetic structure

Abstract: Genetic structure and patterns of genetic variation 

among provenances for eastern white pine (Pinus strobus L.) 

in Quebec were examined by a provenance-progeny test. A 

sample of 66 provenances of eastern white pine from Quebec 

and adjacent Ontario was tested, with each provenance composed of one to eight separate families (a total of 159 families). 

Data were collected on total seedling heights at different ages 

up to 4 years, 4-year branch number, and 3 year bud burst and 

bud set. Analysis of variance showed significant differences 

in each trait among provenances and among families within 

provenances, with one exception. Variance due to provenances 

was on average twice that due to families within provenances. 

All traits were intercorrelated at the provenance level. Two 

principal components summarized 77% of the total variation 

for all traits. Regressions of different traits on geographical 

variables accounted for 16% to 62% of provenance variation 

and disclosed complex patterns of genetic variation among 

provenances. Provenances from the southeastern part of the 

sampling area were taller and burst and set terminal buds later 

than the northwestern ones. The regression models were tested 

using 10-year height and survival from three field tests, which 

gave mixed results. We used the models describing patterns of 

provenance variation to guide seed transfer and delineated two 

provisional breeding zones for eastern white pine in Quebec.

707. Lipow SR, Vance-Borland K, St. Clair JB, 

Henderson JA, McCain C. 2007.In situ gene conservation 

of six conifers in Western Washington and Oregon. Western 

Journal of Applied Forestry 22:176-187.

Type: Journal 

Geographic Area: Pacific Northwest USA

Keywords: gap analysis, biodiversity, genetic resources, conservation planning, GIS

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 251

Link: http://www.invemar.org.co/redcostera1/invemar/

docs/RinconLiterario/2011/febrero/AG_8.pdf

710. Mahalovich MF, Dickerson GA. 2004. Whitebark 

pine genetic restoration program for the Intermountain West 

(United States). In: Sniezko R, Samman S, Schlarbaum SE, 

Kriebel HB, editors. IUFRO Working Party 2.02.15. Fort Collins 

(CO): USDA Forest Service, Rocky Mountain Research Station. 

RMRS-P-32. p 181-187.

Type: Government Document

Geographic Area: Western USA

Keywords: white pine blister rust resistance, fire restoration, 

genetic conservation, seed transfer guidelines

Abstract: A strategy to restore whitebark pine communities is 

presented that emphasizes genetic resistance to white pine blister rust (Cronartium ribicola Fisch.) and mountain pine beetle 

(Dendroctonus ponderosae Hopkins), in combination with an 

active tree planting program. Early and active intervention 

may prevent listing of whitebark pine under the Endangered 

Species Act and further aid in the successful recovery of the 

grizzly bear (Ursus arctos horribilis). The restoration program 

initiated in 2001 includes a multi-State effort (Idaho, Montana, 

Oregon, Nevada, Wyoming, and Washington) designating permanent leave trees, emphasizing clean trees in high blister rust 

areas or areas with a high incidence of mountain pine beetle, or 

areas where both conditions are present. Cone collections from 

these trees will provide an immediate seed source for fire restoration, reforestation, ex situ genetic conservation, and seedlings to be screened for blister rust resistance. Pollen will be 

collected for genetic conservation and to advance blister rust 

resistance in seed and breeding orchards. Data generated from 

the rust screenings will identify whitebark pine seed sources 

that provide high levels of blister rust resistance and provide 

information needed to refine seed transfer guidelines. Leave 

trees elevated to elite-tree status, as identified by their rust 

resistant progeny in the rust screenings, will serve as a seed 

source for operational collections and seed trees for natural 

regeneration. Survivors from the blister rust screening will be 

planted in clone banks for genetic conservation purposes, to 

serve as donors for future seed orchard establishment, and to 

facilitate selective breeding for blister rust resistance.

Link: http://prdp2fs.ess.usda.gov/Internet/FSE_DOCUMENTS/stelprdb5341433.pdf

711. Mahalovich MF, Burr KE, Foushee DL. 2006.

Whitebark pine germination, rust resistance, and cold hardiness 

among seed sources in the Inland Northwest: planting strategies 

for restoration. In: Riley LE, Dumroese RK, Landis TD, editors. 

Forest and Conservation Nursery Associations—2005. Fort 

Collins (CO): USDA Forest Service, Rocky Mountain Research 

Station. Proceedings RMRS-P-43. p 91-101.

Type: Government Document

northward seed transfer within a distance of 2–2.5° in latitude 

could be acceptable to promote stand growth while incurring no 

statistically significant increase in cold damage.

709. Lynch M, Walsh B. 1998. Genetics and analysis of 

quantitative traits. Sunderland (MA): Sinauer Associates, Inc. 

980 p. 

Type: Book

Geographic Area: Global

Compilers’ Keywords: genotype-environment interaction

Abstract: Genetics and Analysis of Quantitative Traits brings 

together the diverse array of theoretical and empirical applications of quantitative genetics under one cover, in a way that is 

both comprehensive and accessible to anyone with a rudimentary understanding of statistics and genetics. What was originally envisioned as a single text has now become two, with the 

focus of this first book being on the basic biology and methods of analysis of quantitative characters. (The second book, 

Evolution and Selection of Quantitative Traits, will address 

the basic principles that govern the evolutionary dynamics of 

quantitative characters under the forces of mutation, random 

genetic drift, and natural and/or artificial selection, integrating the theory of evolution of such characters with existing 

empirical data from natural and domesticated plant and animal 

populations.) Written as a general text in quantitative genetics, 

the book will also be useful as a basic reference for the seasoned professional. Throughout, central theoretical concepts 

are developed from first principles. To aid the less statistically 

sophisticated reader, several chapters and appendices focus on 

the basic statistical tools needed to digest the book. Wherever 

possible, theoretical and analytical concepts are illustrated with 

empirical examples from diverse settings. Three major features 

of Genetics and Analysis of Quantitative Traits distinguish it 

from earlier work. First, it reflects the explosive influx over the 

past few years of quantitative-genetic thinking into evolutionary biology. Second, in animal breeding, enormous strides have 

been made in the development of new techniques for estimating 

breeding values (for the purposes of identifying elite individuals in selection programs) and for estimating variance components from samples of complex pedigrees. In this text’s last two 

chapters, the authors outline the basic principles of complex 

pedigree analysis, without getting bogged down in technical 

details. Third, Genetics and Analysis of Quantitative Traits 

provides a broad overview of the newly emerging array of 

techniques for quantitative-trait loci (QTL) analysis, currently 

one of the most active fields of quantitative-genetic research. 

Genetics and Analysis of Quantitative Traits contains numerous 

fully-worked examples and illustrations of theoretical concepts, 

as well as over 2,000 references with indices by subject, author, 

and organism. In addition, the authors maintain a World Wide 

Web site featuring up-to-date lists of computer programs and 

on-line resources, and added information on various topics presented in the text.

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252 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

analysis for the four Pyrenean plants studied (Trifolium alpinum, Festuca eskia, Festuca gautieri and Rumex scutatus), 

based on random amplified polymorphic DNA (RAPD) 

marker analysis, unweighted pair-group method with arithmetic averages (UPGMA) analysis and Mantel correlograms 

comparing geographic and genetic distances. Results: The 

genetic analysis allowed definition of two main evolutionarily significant units (ESUs) for the plants under study. 

Although the limit between the two zones was slightly variable according to the species considered, an eastern and a 

western ESU was consistently observed. This delineation 

was concordant with the main phytogeographic boundaries of the French Pyrenees. RAPD markers and associated 

Mantel correlograms can be useful to draw ESUs for individual species when the sampling intensity is relatively 

dense, and similarities were revealed between species sharing the same distribution range. This delineation allowed 

integration of intraspecific plant variation in the management of natural resources for revegetation in the Pyrenees. 

Nevertheless, caution is needed for the establishment of seed 

pools in order to maximize genetic diversity in each of the 

pools during collection and production.

713. Merkle SA, Adams WT. 1987. Patterns of 

allozyme variation within and among Douglas-fir breeding 

zones in southwest Oregon. Canadian Journal of Forest 

Research 17:402-407.

Type: Journal 

Geographic Area: Oregon, USA

Compilers’ Keywords: breeding zone, seed transfer, 

Pseudotsuga menziesii, adaptive variation

Abstract: Gametophytes from wind-pollinated seeds of 

Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco var. menziesii) parent trees (mean 56) in each of 22 breeding zones 

in southwest Oregon were analyzed electrophoretically for 

gene frequency patterns at 27 loci. Allozyme variability 

levels were high, as shown by breeding-zone averages for 

the percentage of polymorphic loci (71.7%, 0.99 criterion), 

mean number of alleles per locus (2.46), and expected heterozygosity (0.178). Differences among zones in allele frequency were significant (p < 0.05) for only 2 of the 27 loci 

surveyed, and analysis of hierarchical population structure 

showed that less than 1% of genetic diversity was attributable to differences among breeding zones. Genetic distance 

between zones was small (D = 0.0013) and, in general, bore 

no relation to geographical or environmental distance. The 

limited allozyme differentiation among zones contrasts 

strikingly with the environment-related variation in seedling 

quantitative traits previously reported for southwest Oregon 

Douglas-fir. Allozymes do not appear to be useful for mapping patterns of adaptive variation or for certifying Douglasfir seed in this region.

Geographic Area: Northwestern USA

Keywords: Pinus albicaulis, progeny test, genecology, heritability, electrolyte leakage test, index of injury

Abstract: A synthesis of several studies highlights aboveaverage performing seed sources (n = 108) of whitebark pine 

(Pinus albicaulis), which practitioners can utilize for restoration, wildlife habitat improvement, and operational planting 

programs. It is the first report of this magnitude of blister rust 

resistance for this species. Whitebark pine does have genetic 

variation and demonstrated resistance to white pine blister 

rust, increasing from the southeast to the northwest in the 

Inland Northwest. Early outplanting reports have shown that 

some seedlings have frost damage or exhibit increased mortality in cold pockets or swales. Cold hardiness, measured 

in late winter on a smaller sample of sources (n = 55), also 

showed genetic variability increasing from the northwest to 

the southeast. Seed zones were delineated by Mahalovich 

and Hoff (2000) based on information on relative rust hazard and demarcation of mountain ranges. These geographic 

seed zones support conservative seed transfer with a special 

emphasis on blister rust infection levels. Sufficient variability exists to maintain these seed zone boundaries, because 

whitebark pine exhibits more of an intermediate adaptive 

strategy as compared to the generalist adaptive strategy of 

western white pine (P. monticola). Based on this composite 

information, it is feasible to outplant whitebark pine without the additional delay of waiting until blister rust resistant 

seedlings are developed from a breeding program. There are 

sources within each seed zone that have both rust resistance 

and greater cold hardiness, so those factors should not limit 

tree planting for restoration or critical wildlife habitat improvement objectives. Typical stock orders involve container-grown seedlings. A comparison between Economy and 

copper-lined Ray Leach Super Cell Cone containersTM (10 

in3 [164 cm3]) shows no advantage to using copper lining.

Link: http://www.treesearch.fs.fed.us/pubs/26662

712. Malaval S, Lauga B, Regnault-Roger C, Largier 

G. 2010. Combined definition of seed transfer guidelines for 

ecological restoration in the French Pyrenees. Applied Vegetation 

Science 13:113-124.

Type: Journal 

Geographic Area: France, Europe

Keywords: genetic distance, genetic variation, local provenance, mantel correlograms, native plant, phytogeographic 

boundaries, RAPD markers, seed collection strategy

Abstract: Can genetic tools combined with phytogeography help to define local plants and how geographically close 

the source population should be to the restoration site? The 

main phytogeographic boundaries in the French Pyrenees 

described by different authors were studied and this geographic pattern was compared with the results of genetic 

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Abstract: Prairie restoration is often limited by the availability of appropriate local plant materials. Use of locally adapted 

seed is a goal in restoration, yet little information to inform 

seed transfer guidelines is available for native plant species. 

We established common gardens of five plants (Eriophyllum 

lanatum var. leucophyllum, Epilobium densiflorum, Potentilla 

gracilis var. gracilis, Lupinus polyphyllus var. polyphyllus, and 

Saxifraga oregana) frequently used in prairie restoration in the 

Pacific Northwest of North America to determine if populations differed in morphological and phenological traits and 

whether this variability was structured by geography, climate, 

or habitat. Ordination techniques were used to summarize 

the observed variability of multiple traits for each species. 

Ordination distance was significantly correlated with geographic distance in L. polyphyllus var. polyphyllus, and populations of this species differed significantly among geographic 

groups within an ecoregion. Little or no spatial structure was 

detected in the remaining species, despite correlations between 

ordination scores and monthly temperatures. We suggest that 

a single seed zone in the Willamette Valley ecoregion may be 

appropriate for all species examined except L. polyphyllus var. 

polyphyllus. Ecoregions in general may be useful boundaries 

for seed transfer zones, especially in regions with relatively 

little topographical or climatic variation. 

Link: http://www.snprsip.com/resources/publications/seedplant-development/Miller_2011.pdf

716. Montalvo AM, Ellstrand NC. 2001. Nonlocal 

transplantation and outbreeding depression in the subshrub Lotus 

scoparius (Fabaceae). American Journal of Botany 88:258-269.

Type: Journal 

Geographic Area: California, USA

Keywords: coastal sage scrub, common garden, cumulative 

fitness, ecological genetics, environmental distance, geographic 

distance, genetic distance, local adaptation, restoration, seed 

source

Abstract: The genetic background of transplants used to 

create or augment wild populations may affect the long-term 

success of restored populations. If seed sources are from differently adapted populations, then the relative performance of 

progeny from crosses among populations may decrease with 

an increase in genetic differences of parents and in the differences of parental environments to the transplant location. We 

evaluated the potential for such outbreeding depression by hybridizing individuals from six different populations of Lotus 

scoparius var. scoparius and L. s. var. brevialatus. We used allozyme data to calculate genetic distances between source populations, and compiled climatic data and measured soil traits 

to estimate environmental distances between source populations. We found significant outbreeding depression following 

controlled crosses. In the greenhouse, the success of crosses 

(seeds/flower × seedlings/seed) decreased with increasing 

714. Michalski SG, Durka W. 2012. Assessment of 

provenance delineation by genetic differentiation patterns 

and estimates of gene flow in the common grassland plant 

Geranium pratense. Conservation Genetics 13:581-592.

Type: Journal 

Geographic Area: Germany, Europe

Keywords: Geranium pratense, population structure, local 

provenance, AFLP, outcrossing rate, genome scan

Abstract: The use of local provenances in restoration, agriculture and forestry has been identified and widely implemented 

as a measure for species and community conservation. In practice, provenances are often delineated based on climate, geomorphology and correlated spatial descriptors following the 

boundaries of larger natural regions. They are thought to comprise genetically homogenous plant material. Here we investigate genetic variation at AFLP loci in 26 natural populations of 

the regionally common grassland species Geranium pratense, 

which is often used in seed mixtures. Most studied populations are located in one previously delineated provenance in 

Germany. We assess within- and among provenance differentiation patterns and aspects of gene flow by investigating the 

mating system, the genetic structure at regional and local scale, 

gene dispersal and potential selective mechanisms that may 

have contributed to differentiation patterns found. Compared 

to other herbaceous, insect-pollinated grassland species and 

despite being outcrossed (mean tm = 0.88), G. pratense showed 

low genetic diversity (mean HE = 0.15), considerable genetic 

differentiation among populations within provenances (mean 

pairwise FST = 0.20) and a pronounced within-population 

spatial genetic structure (mean Sp = 0.064). A genome scan 

approach identified three potentially adaptive loci. However, 

their allelic frequencies were only weakly related to climatic 

parameters thus providing little evidence for adaptive divergence. Nevertheless, the distribution of genetic diversity and 

derived gene dispersal estimates indicate limited dispersal 

ability, suggesting that gene flow at distances larger than 10 

km is negligible. Our findings may question the approach of 

delineating provenances by general criteria, and highlight the 

importance of species specific studies on differentiation and 

adaptation patterns.

Link: http://www.ufz.de/export/data/global/33126_Michalski_Durka_2012.pdf

715. Miller SA, Bartow A, Gisler M, Ward K, Young 

AS, Kaye TN. 2011. Can an ecoregion serve as a seed transfer 

zone? Evidence from a common garden study with five native 

species. Restoration Ecology 19:268-276.

Type: Journal 

Geographic Area: Pacific Northwest USA

Keywords: common garden, ecoregion, restoration, seed transfer zone

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254 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

ranging from 72% to 100% for individual provenances, but 

many Florida and Gulf coast populations exhibited reduced 

survival and winter hardiness in New Jersey, with 44–94% 

survival and 11–67% of surviving trees experiencing winter 

dieback. Height growth varied significantly between planting sites and provenances, and a significant site by provenance interaction suggested that relative performance of 

provenances varied across sites. All populations grew significantly more at Hofmann Forest, North Carolina than at the 

two New Jersey plantations, with mean 2-year height growth 

ranging from 54 to 96 cm for individual provenances at this 

site. By comparison, mean height growth ranged from 23 to 

63 cm at Richard Stockton College, and from 12 to 32 cm 

at Brendan T. Byrne State Forest, New Jersey. Within individual sites, we identified polynomial relationships between 

height growth and latitude of origin, with southern Atlantic 

coastal populations among the best performers at all sites. 

Patterns of growth phenology were significantly correlated 

with latitude of origin. Spring growth rate and the percent 

of total growth occurring during spring were positively correlated with latitude, while fall growth rate and the percent 

of growth occurring during fall were negatively correlated 

with latitude. These early results suggest that significant 

provenance variation exists among white-cedar populations, 

and that local sources should be preferred for regeneration 

and restoration purposes.

718. Namkoong G, Conkle MT. 1976. Time trends in 

genetic control of height growth in ponderosa pine. Forest Science 

22:2-12.

Type: Journal 

Geographic Area: California, USA

Keywords: Pinus ponderosa, geographic variation, elevation effects, tree growth, genetic variation

Abstract: Height growth in ponderosa pine plantation with 

71 families in 7 elevational zone sources was analyzed at 

ages 3, 5, 7, 8, 12, 20, 25, and 29. Height growth varied by 

seed source zone and by family within zone. Zonal variances 

were well established at an early age, and the differences were 

maintained with minor variations through the study period. 

Family variances were smaller than zonal and environmental 

variances, the differences were not well correlated between the 

earlier and later phases, and the correlations were sometimes 

negative. Three observed growth phases, related to ecological 

dominance and competition, affected the levels of variance attributable to genetics and environment.

719. Nanos N, González-Martínez SC, Bravo F. 

2004. Studying within-stand structure and dynamics with 

geostatistical and molecular marker tools. Forest Ecology and 

Management 189:223-240.

Type: Journal 

genetic distance between populations revealing genetically 

based outbreeding depression unrelated to local adaptation. 

After outplanting to one native site (in situ common garden), 

field cumulative fitness of progeny (survival × fruit production) 

decreased significantly with mean environmental distance of 

the parental populations to the transplant site, but not with genetic distance between the crossed populations. This result is 

consistent with a disruption of local adaptation. At the second, 

ecologically contrasting common garden, where low survival 

reduced statistical power, field cumulative fitness (survival × 

progeny height) did not decrease significantly with either environmental distance or genetic distance. Overall, intervariety 

crosses were 40 and 50% as fit (seeds/flower × seedlings/seed 

× survival × fruits at the first garden or × height at the second) 

as intravariety crosses. These results suggest that the cumulative outbreeding depression was caused by a combination of 

genetically based ecological differences among populations 

and other genomic coadaptation. We conclude that mixing 

genetically differentiated seed sources of Lotus scoparius may 

significantly lower the fitness of augmented or restored populations. Genetic and environmental similarities of source populations relative to the transplant site should be considered when 

choosing source materials, a practice recommended by recent 

seed transfer policies. Geographic separation was not a good 

surrogate for either of these measures.

717. Mylecraine KA, Kuser JE, Zimmermann GL, 

Smouse PE. 2005. Rangewide provenance variation in 

Atlantic white-cedar (Chamaecyparis thyoides): early survival and 

growth in New Jersey and North Carolina plantations. Forest 

Ecology and Management 216:91-104.

Type: Journal 

Geographic Area: USA

Keywords: Cupressaceae, local adaptation, phenotypic variation, Chamaecyparis thyoides, growth phenology, geographic 

variation

Abstract: Atlantic white-cedar, Chamaecyparis thyoides

(L.) B.S.P., is an important wetland tree species occurring 

along the Atlantic coast of the United States, from Maine 

to Florida, and westward along the Gulf of Mexico coast 

to Mississippi. Within this range, its distribution is patchy 

and disjunct, due to the scattered occurrence of suitable wetland habitat, natural range disjunctions and historic declines. 

Much recent interest has focused on the species, its management and its restoration, but very little is known about 

its geographic pattern of phenotypic variation. This study 

presents second year results from the first rangewide whitecedar provenance test. Rooted cuttings were propagated 

from 34 source populations, encompassing the entire latitudinal range (29°12’–44°20’ N) of the species, and planted 

in three common garden plantations, two in New Jersey 

and one in North Carolina. Survival was generally high for 

Atlantic coastal populations at all three planting locations, 

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with strategies to promote the benefits of different agroforestry 

practices. There is also a requirement for conservation planners to consider that trees found outside protected areas have 

a role in the conservation. In wide-ranging species, promoting 

international collaboration for collecting, characterizing, and 

conserving genetic resources is vital as international consensus 

is needed to streamline phytosanitary procedures, to facilitate 

the exchange of forest reproductive material, and assist genetic 

resource conservation efforts.

Link: https://www.google.com/url?sa=t&rct=j&q=&esrc=s

&source=web&cd=1&ved=0CCoQFjAA&url=http%3A%2

F%2Fwww.iufro.org%2Fdownload%2Ffile%2F5907%2F466

8%2F369-384_pdf%2F&ei=PwZ4Uv2gMaewjAKN34DACg

&usg=AFQjCNFzp_g0nqmcx_F2XFqLA7zTAJYlSA&sig2=

DEsdQQrwOnk-w_-fELAbNQ&bvm=bv.55980276,d.cGE

721. O’Brien EK, Mazanec RA, Krauss SL. 2007.

Provenance variation of ecologically important traits of forest 

trees: implications for restoration. Journal of Applied Ecology 

44:583-593.

Type: Journal 

Geographic Area: Australia

Keywords: ecological restoration, forest trees, genetic variation, jarrah, provenance trial, quantitative traits, seed collection 

zones

Abstract: The source of seed or plant material can have profound implications for the success of restoration efforts because most species exhibit adaptive genetic variation within 

their range. Understanding the geographical distribution of 

ecologically relevant genetic variation and the environmental 

factors driving adaptive divergence within species will help to 

ensure appropriate sourcing of material for ecological restoration. We present a study of geographical variation of ecologically important traits of the forest tree jarrah Eucalyptus 

marginata from a 15-year-old provenance trial in southwestern 

Australia. We assessed trait variation in association with rainfall, latitude and slope position at the site of origin. Survival 

and stem diameter varied at the largest scale, between northern 

and southern jarrah forest provenances. Stem diameter also 

varied among rainfall zones, while latitude was a more important determinant of variation of reproductive traits (flowers 

and buds). None of the environmental variables accounted for 

significant variation of height, growth form or the presence of 

capsules. Slope position at the site of origin did not account for 

significant variation of any trait. Trees from low rainfall sites 

had smaller stem diameters, possibly reflecting selection for 

slower growth. Such a strategy could prevent drought stress 

and may explain why trees from the high rainfall southern jarrah forest, which showed the fastest growth, had the poorest 

survival at the drier northern trial site. Variation in the presence 

of buds and flowers among latitudinal divisions may be because of variation in flowering time, which has been observed 

Geographic Area: Spain, Europe

Keywords: geostatistics, spatial structure, stand dynamics, 

competition, molecular markers

Abstract: We established a circular experimental plot (radius 

= 100 m) that consists of 380 trees of maritime pine (Pinus 

pinaster Ait.). The plot is located in central Spain (Segovia 

province) and within a pure, even aged and naturally regenerated stand of the species. Every tree was initially mapped using 

polar coordinates. The objective of the study was to explore 

the spatial arrangement of some stand attributes such as age of 

trees, diameter at breast height (DBH), total height, diameter 

increment and number of female strobili and cones. Genetic 

data (three highly polymorphic nuclear microsatellites) were 

used to investigate extend of relatedness within the stand and 

differentiate between genetic and environmental causes of 

spatial aggregation. We employed, in the analysis of tree attributes, only geostatistical tools (direct and cross variograms 

as well as their indicator counterparts). The results revealed 

that all the studied attributes are spatially correlated and some 

of them cross-correlated. Competition and height were found 

to be key-variables in stand dynamics, which influence the rest 

of the attributes. Fine-scale genetic structure exists within the 

stand (0–30 m range). Relatedness between neighboring trees 

can partially explain the spatial aggregation of individual tree 

attributes. Geostatistical tools are recommended for this type 

of spatial analysis of forest stands. Their use can provide useful insights into the nature of stand organization and can guide 

future silvicultural treatment toward higher stand productivity 

or better and faster regeneration.

720. Navarro C, Boshier D, Cavers S, Lowe A. 2010.

Genetic resources and conservation of mahogany in Mesoamerica.

Forests and Society 20:369-383.

Type: Journal 

Geographic Area: Mexico

Keywords: mahogany, Swietenia, conservation, management, 

policies, genetic diversity, genetic resource, agroforestry

Abstract: We review here the scale of neutral and adaptive 

genetic variation of mahogany (Swietenia spp.), one of the 

most valuable tree species in the world, and relate this information directly to provenance performance, and seed transfer 

and sourcing recommendations. Further, we explore a series 

of scientific issues directly related to critical day-to-day management of mahogany, giving guidelines to maximise genetic 

diversity outcomes in natural, remnant, sustainably harvested, 

and/or restored forest landscapes. The study concludes that 

conservation and sustainable management of mahogany genetic resources cannot be simply defined in terms of one-sizefits-all solutions, and requires local community involvement to 

prevent illegal logging and ensure mutual benefit. Landscape 

level strategies for the effective management of mahogany 

trees outside of forests urgently need to be developed, together 

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Compilers’ Keywords: Pinus contorta, long-shoot bud development, axillary bud, terminal apical

Abstract: Long-shoot bud development, shoot growth, and 

foliage production were studied in seven provenances of 

Pinus contorta Dougl. ssp. latifolia Engelm. from the major 

sites in British Columbia and one Yukon source growing in a 

provenance trial at Prince George, B.C. Branch terminal apical mitotic activity began in early March and continued until late September. Initiation of axillary buds began in May, 

about 2 weeks after the initiation of the subtending cataphyll. 

Differentiation of dwarf shoots began in early July to mid-August and continued until late October in some sources. Distal 

axillary buds had not always differentiated by late October 

in the southern sources. The duration of the period of apical 

growth and apical size during activity were related to final cataphyll numbers. Provenances with the widest, flattest, dormant 

apices produced the most cataphylls. The two northern provenances had more terminal sterile cataphylls but fewer sterile 

cataphylls lower in the long-shoot bud and shorter mean stem 

unit lengths than the others. Differences among provenances 

in shoot length were due mostly to variation in stem unit numbers. The large proportion of polycyclic long shoots in some 

provenances contributed to variation in dwarf shoot numbers. 

724. O’Reilly C, Owens JN. 1989. Polycyclic growth and 

branching in the upper crown in provenances of lodgepole pine.

Canadian Journal of Forest Research 19:79-87.

Type: Journal 

Geographic Area: Western Canada

Compilers’ Keywords: provenance study, branch numbers, 

Pinus contorta, seed source

Abstract: The percentage of shoots that originated from 

polycyclic growth in first-order branches and branch numbers 

in upper annual segments of leader growth were determined 

from trees of seven provenances of Pinus contorta Dougl. ssp. 

latifolia Engelm. from British Columbia and the Yukon, growing in a provenance trial at Prince George, B.C. The number 

of first-order branches varied among provenances and year of 

growth and was related to the frequency of extra cycles. The 

percentage of polycyclic shoots was greatest in the 2nd and 

4th years of branch growth, although this varied with year and 

provenance. Total branch numbers were influenced mostly by 

variation in first- and second-order branch numbers. Secondorder branch numbers were greatest in the second and third 

segments of leader growth (from the top down), but this was 

influenced by year of growth and provenance. The high levels of second-order branching corresponded to the greater 

frequency of polycyclic branches in these positions. The fastgrowing southern provenance had the most branches, while 

the slow-growing Yukon source had the least. The slowergrowing provenances had the most first-order branches per 

unit height. 

previously among E. marginata populations. However, variation among replicate blocks within the trial suggests that the 

environment also strongly influences expression of these traits. 

We have demonstrated divergence of several ecologically 

important traits in association with different types of environmental variation. Our findings support an argument for ‘habitat matching’ when sourcing material for restoration; however, 

differences among trait types in the distribution of variation 

highlight the need to consider environmental variation at a 

range of geographical scales. Consideration of ecologically 

important genetic variation within species is important and this 

information should be integrated into seed collection strategies 

for ecological restoration.

722. O’Neill GA, Nigh G, Wang T, Ott PK. 2007.

Growth response functions improved by accounting for 

nonclimatic site effects. Canadian Journal of Forest Research 

37:2724-2730.

Type: Journal 

Geographic Area: Western Canada

Compilers’ Keywords: lodgepole pine, Pinus contorta, provenance study

Abstract: Growth response functions (GRFs) that relate the 

growth of a population to the climate of the sites in which 

it is tested are gaining attention for their ability to predict 

impacts of climate change on tree growth. However, nonclimatic site to site variation introduces error into GRFs. Using 

data from a large lodgepole pine (Pinus contorta Dougl. ex 

Loud.) provenance test in British Columbia and the Yukon, 

Canada, a technique is presented that accounts for the effect 

of non-climatic variation in GRFs. The mean height of the 

“local” provenances at each test site was used to predict “site 

height” from site climate variables in multiple regression. 

Residuals from the site height equation provided an index of 

the nonclimatic effect for each site and were included as a 

covariate in quadratic GRFs that related provenance height 

at each test site to mean annual temperature at each test site. 

Inclusion of the nonclimatic index in the model resulted in 

a moderate or large displacement of GRFs for 25% of the 

provenances, while increasing mean R2 values for 138 of 140 

provenances and decreasing the root mean squared error for 

113 of 140 provenances. These results suggest that inclusion 

of the nonclimatic index in GRF models could substantially 

affect height predictions for some provenances and reduce 

prediction error for most provenances.

723. O’Reilly C, Owens JN. 1987. Long-shoot bud 

development, shoot growth, and foliage production in provenances 

of lodgepole pine. Canadian Journal of Forest Research 

17:1421-1433.

Type: Journal 

Geographic Area: Western Canada

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approach has considerable potential to help prevent decreased 

growth and yield due to the planting of maladapted seed.

727. Parker WH, van Niejenhuis A. 1996.

Regression-based focal point seed zones for Picea mariana 

from northwestern Ontario. Canadian Journal of Botany 

74:1227-1235.

Type: Journal 

Geographic Area: Ontario, Canada

Keywords: black spruce, Picea mariana, adaptive variation, focal point seed zones 

Abstract: The results of a recent study of adaptive variation 

of black spruce in northwestern Ontario, together with additional freezing damage data, were used to produce regressionbased focal point seed zones for this species. The procedure 

required two data bases as follows: (i) the biological data derived from two common garden growth trials, one greenhouse 

trial, and freezing trials of 75 black spruce seed sources and (ii) 

climatic data for the period 1951-1980. Principal components 

analysis (PCA) was used to summarize the main components 

of growth and freezing variation, and the PCA axis scores for 

the seed sources were regressed against climatic variables. The 

regression equations were used to model the patterns of adaptive variation, and these patterns were graphically reproduced 

as contour maps by a geographic information system (GIS). 

A series of focal point seed zone maps for black spruce was 

produced by GIS intersection of the regression-based contour maps. Focal point seed zones were more restricted in the 

south near Lake Superior, reflecting the more rapidly changing climate in this part of the study area. Since black spruce 

is closely adapted to local climate, these results will be useful 

to formulate successful seed transfers in this area. There are 

additional potential applications for matching seed sources 

to changing climates and for the identification of genetically 

unique populations. 

Link: http://flash.lakeheadu.ca/~whparker/FPSZ1996reprint.pdf

728. Parsons MC, Jones TA, Larson SR, Mott IW, 

Monaco TA. 2011. Ecotypic variation in Elymus elymoides

subsp. brevifolius in the northern Intermountain West. Rangeland 

Ecology & Management 64:649-658.

Type: Journal 

Geographic Area: Western USA

Keywords: AFLP, common garden, Great Basin, ecotype, 

squirreltail

Abstract: Bottlebrush squirreltail (Elymus elymoides [Raf.] 

Swezey) is an important native bunchgrass for rangeland restoration in western North America. This species is taxonomically 

complex and has diverged into as many as four subspecies, 

including subsp. brevifolius, for which four geographically 

725. Oswald BP, Neuenschwander LF. 1995.

Mortality of western larch seedlings in relation to seedbed 

characteristics at the dry end of its ecological range. Bulletin of 

the Torrey Botanical Club 122:101-108.

Type: Journal

Geographic Area: Idaho, USA

Keywords: safe sites, western larch, seedbed, seedling mortality, Larix

Abstract: The effects of different seedbeds on western larch 

(Larix occidentalis Nutt.) germination and seedling mortality 

were assessed in a grand fir/ninebark (Abies grandis ((Dougl. 

ex. D. Don) Lindl.))/ (Physocarpus malvaceus ((Greene) 

Kuntze)) habitat type in northern Idaho, U.S.A. Two plots were 

established on each of four sites, and 150 western larch seeds 

(66% viable) were distributed on each of 16 randomly applied 

seedbed modification combinations (burned or mechanical 

scarified, mineral soil or duff, shaded or non-shaded, caged or 

non-caged for protection from predation). Germination percentages and seedling development were significantly greater 

(4–10 times) on caged units than on uncaged units by midJuly. Shading did not significantly affect germination percentage or initial seedling establishment. No significant differences 

occurred in germination or initial seedling establishment between burned and unburned treatments. Mortality of seedlings, 

most likely caused by a month-long dry period with high 

temperatures, resulted in no significant differences in seedling 

numbers among treatments by mid-August the first year after 

seed dispersal. On the dry end of this species’ range, there does 

not appear to be any difference in seedling survival from seedbed modification.

726. Parker WH. 1992. Focal point seed zones: site-specific 

seed zone delineation using geographic information systems. 

Canadian Journal of Forest Research 22:267-271.

Type: Journal 

Geographic Area: Ontario, Canada

Compilers’ Keywords: Jack pine, Pinus banksiana, GIS, provisional zones, seed transfer zone, common garden

Abstract: A new site-specific approach to defining seed zones 

in North American conifers is described. Using focal point 

seed zones, an individual site to be reforested becomes the focal point, and a unique seed zone is established for that site 

as needed. This approach depends upon (i) obtaining good 

comparative data in adaptive characteristics from throughout 

the range to be regenerated based upon a series of short-term 

growth tests in a common garden and (or) greenhouse and 

(ii) graphic analysis of multivariate summary scores by geographic information systems software to delimit boundaries of 

unique seed zones for any location to be reforested. A sample 

focal point seed zone is delineated for jack pine (Pinus banksiana Lamb.) reforestation of a site in northern Ontario. This 

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730. Pedersen AP, Hansen JK, Mtika JM, Msangi 

TH. 2007. Growth, stem quality and age-age correlations in a 

teak provenance trial in Tanzania. Silvae Genetica 56:142-148.

Type: Journal 

Geographic Area: Tanzania, Africa

Keywords: Tectona grandis, land races, provenance variation

Abstract: This study examines the growth and stem straightness variation between teak (Tectona grandis L. f.) provenances 

in a 30-year-old field trial in the coastal lowland of Tanzania. 

The results are compared with earlier results. The findings 

demonstrate that it is possible to find large height and yield 

differences between teak provenances. The best performing 

provenance grows some 10% (height) to 40% (volume) better 

than the average. The provenance Topslip, India, consistently 

proved to be outstanding in terms of growth and stem quality 

confirming earlier evaluations at age 5 and 17 years. Tanzanian 

land races also proved to be superior, especially as regards 

stem straightness.

Link: http://www.silvaegenetica.com/fileadmin/content/dokument/archiv/silvaegenetica/56_2007/56-3-4-142.pdf

731. Perks MP, McKay HM. 1997. Morphological and 

physiological differences in Scots pine seedlings of six seed origins.

Forestry 70:223-232.

Type: Journal 

Geographic Area: Scotland, United Kingdom

Compilers’ Keywords: Pinus sylvestris, provenance study

Summary: One-year-old seedlings of Scots pine (Pinus sylvestris L.) of four native seed origins (Loch Maree Islands, 

Glengarry/Glen Morriston, Glen Affric and Abernethy), a commercial British seedlot, and a seedlot from Hedesunda, in middle Sweden, were compared at monthly intervals from October 

1993 to April 1994. Seedling morphology, root condition, root 

frost hardiness and bud dry matter were determined at each 

date. There were clear morphological differences among seed 

origins. Seedlings raised from the commercial seedlot (A70) 

were larger but had a poorer root : shoot ratio than the other 

seed origins. Of the native pines tested, the Loch Maree Islands 

origin allocated a larger proportion of its photosynthate to fine 

roots and needles and smaller proportion to woody structures. 

Seedlings raised from the commercial British seedlot tended to 

have poorer bud lignification than the other origins and also, 

in autumn, higher electrolyte leakage rates from its fine roots. 

During winter, the Swedish origin had the lowest fine root 

electrolyte leakage. Seedlings of all origins showed a progressive increase in fine root hardiness towards mid-winter with 

maximum hardiness (-7 °C) in January. Dehardening occurred 

over subsequent months reaching -3 °C in April. Differences 

among origins were evident. The Swedish seedlot developed 

greater frost resistance than the other origins, hardening began 

earlier in autumn and dehardening began later in spring. The 

distinct races have been described (A, B, C, and D). Of these four 

races, only C occurs in the northern Intermountain West. Our 

objectives were to describe phenotypic and genetic variation 

within C and to ascertain its taxonomic status. We evaluated 32 

populations of C collected across the northern Intermountain 

West for a battery of biomass, phenological, and functional 

traits in common-garden settings in the field and greenhouse. 

Genetic variation was assessed with the use of amplified fragment length polymorphism (AFLP) markers, and correlations 

were calculated among phenotypic, genetic, environmental, 

and geographic distance matrices with the use of Mantel tests. 

Values for these four distance measures were positively correlated, suggesting that environmental heterogeneity and isolation by distance are shaping ecotypic divergence driven by 

natural selection. We describe three phenotypic zones for C 

that correspond to previously established ecoregion boundaries. Because genetic data group C apart from subsp. brevifolius

races A, B, and D, which originate in the Rocky Mountains and 

western Great Plains, the so called race C merits description as 

a new subspecies apart from subsp. brevifolius.

729. Pastorino MJ, Ghirardi S, Grosfeld J, Gallo LA, 

Puntieri JG. 2010. Genetic variation in architectural seedling 

traits of Patagonian cypress natural populations from the extremes 

of a precipitation range. Annals of Forest Science 67:508-508.

Type: Journal 

Geographic Area: Argentina, South America

Keywords: differentiation, phenotypic plasticity, heritability, 

Austrocedrus

Abstract: Species distributed along wide environmental ranges are expected to be either plastic or adapted to local optima. 

The elucidation of which of these alternatives prevails, is crucial in planning breeding and conservation strategies for not yet 

domesticated species. Austrocedrus chilensis (Cupressaceae) is 

the most commercially important conifer of the temperate forests of Argentina and the target of a domestication program. 

A steep precipitation gradient characterizes its Argentinean 

range. Variation within and differentiation among four natural 

populations of this Patagonian cypress representative of two 

contrasting precipitation regimes (> 1,300 and 330 mm per y) 

were assessed by analyzing several morpho/architectural traits 

in one-year-old seedlings grown in a greenhouse. Progenies 

from one of the two xeric populations did not differ from those 

corresponding to both humid-site populations. The two most 

variable populations in terms of additive genetic variance belonged to contrasting precipitation regimes. Differentiation 

among populations was low as measured by each and every 

variable (average Qst = 0.088). Morphological traits related to 

the main axis size would have a dubious adaptive meaning. 

The results suggest that the Patagonian cypress would have not 

evolved genetic pools adapted to local optima, and thus it appears to be a phenotypically plastic species, at least regarding 

growth at juvenile age.

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variation. Individual plants from each of the 12 sites were genotyped using six AFLP primer combinations detecting DNA 

variation within and among all three species. Genetic differences between species were high enough to render comparisons among species impractical, so each species was analyzed 

separately for differences between populations and variability 

within populations. Similarity coefficients were significantly 

greater within collection sites versus among collection sites 

indicating divergence between populations. Within-population 

genetic diversity was not correlated with elevation for any 

of the three species. Analysis of molecular variance revealed 

that 66% (A. acuminatum), 83% (A. passeyi), and 64% (A. 

brandegei) of observed variation is found within populations. 

Genetic divergence among populations (φST) was higher in 

the widely distributed species, suggesting that interpopulation 

gene flow may be negatively correlated with range size. Allium 

acuminatum and A. brandegei individuals cluster into groups 

corresponding strictly to collection sites based on neighborjoining analysis of the total number of DNA polymorphisms 

between individual plants. Allium passeyi populations, however, had less overall genetic variation between populations. 

Genetic isolation by distance appeared responsible for much 

of the variability among populations, although there was one 

notable exception showing significant differences between two 

geographically close populations in A. acuminatum.

734. Raddad EAY. 2007. Ecophysiological and genetic 

variation in seedling traits and in first-year field performance of 

eight Acacia senegal provenances in the Blue Nile, Sudan. New 

Forests 34:207-222.

Type: Journal 

Geographic Area: Sudan, North Africa

Keywords: agroforestry, adaptation, clay plain, gum belt, reforestation, relative growth rate

Abstract: The genetic variation in seed weight, seed number 

per kg and seedling traits was compared among eight Acacia 

senegal provenances originating from the clay plain (east) and 

sand plains (west) of the gum belt in Sudan. The main objective 

of this study was to identify germplasm sources of A. senegal

that have a good seed germination capacity and seedling traits 

suitable for reforestation in the clay-soil part of the dryland 

gum belt in the Blue Nile region in Sudan. A specific objective 

was to tentatively explore the adaptive strategy of A. senegal

populations. Seventeen-week old seedlings were planted in the 

field at spacing of 3 m · 3 m, giving 100 trees per plot and replication; within a randomized complete block design with four 

replications. The experimental site was in the clay plain region. Seed variables showed significant differences. Clay plain 

provenances showed considerable variation in seed weight and 

seed number. They had the smallest seed weight but the highest seed number, while the sand (western) provenances had the 

largest seed weight but lowest seed number. Seedling branch 

commercial seedlot hardened later than the other origins but 

reached a similar level of frost hardiness by January. Of the 

native pines, seedlings of the Loch Maree Islands origin were 

slowest to develop root hardiness.

Link: http://forestry.oxfordjournals.org/content/70/3/223.full.

pdf

732. Pharis RP, Ferrell WK. 1966. Differences in drought 

resistance between coastal and inland sources of Douglas fir.

Canadian Journal of Botany 44:1651-1659.

Type: Journal 

Geographic Area: Western North America

Compilers’ Keywords: Pseudotsuga menziesii, provenance 

study

Abstract: By two drought-hardiness tests, “time to death” and 

“soil-moisture content at the death point,” Douglas-fir seedlings from three coastal sources were shown to be less drought 

resistant than those from five inland sources. Lethal needlemoisture values, useful as an index of whole plant viability, 

were established for the various sources. Two of the coastal 

sources differed from five inland sources in the level of these 

values, but the lethal points for two other sources from the 

Oregon Cascade Range were similar to the five inland sources. 

Needle moisture appears to be a workable index for determining the whole plant viability except when the plant is very close 

to its time of death. Plants could also be classified into coastal 

and inland groups on the basis of their needle moisture under 

well-watered conditions, with the exception of seedlings from 

the Arizona source which are like the coastal group. 

733. Phillips NC, Larson SR, Drost DT. 2008.

Detection of genetic variation in wild populations of three 

Allium species using amplified fragment length polymorphisms.

HortScience 43:637-643.

Type: Journal 

Geographic Area: Utah, USA

Compilers’ Keywords: genetic diversity, germplasm, divergence, AFLP, geographic variation, genetic isolation, native 

onion

Abstract: Three wild onion species native to the intermountain west in the United States—Allium acuminatum, A. brandegei, and A. passeyi—show horticultural potential, but little 

is known about patterns of genetic diversity among localized 

populations and geographical regions. We examined amplified 

fragment length polymorphisms (AFLP) within and among 

five Allium acuminatum, four A. brandegei and three A. passeyi

collection sites in Utah. These three congeners with contrasting abundance and distribution patterns provide an opportunity 

to investigate the role of geographic distance, altitude, and rarity in patterns of genetic divergence. The collection sites were 

selected along an altitudinal gradient to reflect ecogeographic 

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Link: http://www.ncbi.nlm.nih.gov/pubmed/19732337

736. Rehfeldt GE. 1979. Ecological adaptations in Douglasfir (Pseudotsuga menziesii var. glauca) populations I. North Idaho 

and north-east Washington. Heredity 43:383-397.

Type: Journal 

Geographic Area: Pacific Northwest USA

Compilers’ Keywords: provenance, genetic × environment 

interaction, seed transfer guidelines, tree growth, elevation

Summary: Growth, phenology and frost tolerance of seedlings from 50 populations of Douglas-fir (Pseudotsuga menziesii var. glauca) were compared in 12 environments. Statistical 

analyses of six variables (bud burst, bud set, 3-year height, 

spring and fall frost injuries, and deviation from regression of 

3-year height on 2-year height) showed that populations not 

only differed in mean performance, but also reacted differently 

to the environmental gradient. Most of the population environment interaction was attributable to heterogeneous regressions of population means on environmental means. For all 

variables except growth rate, the variance of heterogeneous regression coefficients was explained by convergence of regression lines to a common point on the environmental gradient. 

Consequently, mean values for populations were significantly 

correlated with regression coefficients. Thus, main effects 

of populations in those single environments that induced the 

greatest mean differences reflected the interaction. Multiple regression analyses associated adaptive differentiation of populations with geographic and ecologic characteristics of the seed 

source. Differentiation was controlled primarily by elevation 

and secondarily by latitude. Whereas longitude was a minor 

factor, habitat types accounted for no differentiation beyond 

that associated with elevation, a factor closely correlated with 

habitat types. From these results it is recommended that seed 

for reforestation should not be moved more than 140 m elevation, l6° latitude, or 27° longitude in northern Idaho and 

eastern Washington.

Link: http://www.treesearch.fs.fed.us/pubs/38633

737. Rehfeldt GE. 1982. Ecological adaptations in Douglasfir populations. II. Western Montana. Ogden (UT): USDA Forest 

Service, Intermountain Forest and Range Experiment Station. 

RP-INT-295. 12 p.

Type: Government Document

Geographic Area: Montana, USA

Compilers’ Keywords: Pseudotsuga menziesii var. glauca, 

common garden, provenance study, tree populations, seed 

transfer guidelines

Research Summary: Seedlings from 50 populations of 

Douglas-fir (Pseudotsuga menziesii var. glauca) from Montana 

west of the Continental Divide were compared in nursery, laboratory, and shadehouse studies. Analyses of variance indicated 

number, root length, root to shoot ratio and shoot dry weight 

differed significantly among the provenances 12 weeks after 

germination. Clay provenances had the highest branch number 

and shoot dry weight but the shortest roots and lowest root to 

shoot ratio. This was interpreted as showing better adaptation 

to the site in these local provenances in comparison to those 

originating from the western sandy soil regions. High positive 

correlations were observed between seedling variables, such 

as root nodule and branch numbers; this could be used for 

early selection. The variation was greater between provenance 

groups than within them, suggesting that especially selection 

among groups would yield genetic gain.

735. Ramirez-Valiente JA, Lorenzo Z, Soto A, 

Valladares F, Gil L, Aranda I. 2009. Elucidating the role 

of genetic drift and natural selection in cork oak differentiation 

regarding drought tolerance. Molecular Ecology 18:3803-3815.

Type: Journal 

Geographic Area: Spain, Europe

Keywords: adaptation, carbon isotope discrimination, drift, 

FST, leaf size, QST, Quercus suber, selection

Abstract: Drought is the main selection agent in Mediterranean 

ecosystems and it has been suggested as an important evolutionary force responsible for population diversification in these 

types of environments. However, population divergence in 

quantitative traits can be driven by either natural selection, 

genetic drift or both. To investigate the roles of these forces 

on among-population divergence in ecophysiological traits 

related to drought tolerance (carbon isotope discrimination, 

specific leaf area, leaf size and leaf nitrogen content), we 

compared molecular and quantitative genetic differentiation 

in a common garden experiment including thirteen cork oak 

(Quercus suber L.) populations across a gradient of rainfall and 

temperature. Population differentiation for height, specific leaf 

area, leaf size and nitrogen leaf content measured during a dry 

year far exceeded the molecular differentiation measured by 

six nuclear microsatellites. Populations from dry-cool sites 

showed the lowest nitrogen leaf content and the smallest and 

thickest leaves contrasting with those from humid-warm sites. 

These results suggest (i) these traits are subjected to divergence 

selection and (ii) the genetic differences among populations are 

partly due to climate adaptation. By contrast, the low amongpopulation divergence found in basal diameter, annual growth 

and carbon isotopic discrimination (a surrogate for water use 

efficiency) suggests low or no divergence selection for these 

traits. Among-population differentiation for neutral markers 

was not a good predictor for differentiation regarding the quantitative traits studied here, except for leaf size. The correlation 

observed between the genetic differentiation for leaf size and 

that for molecular markers was exclusively due to the association between leaf size and the microsatellite QpZAG46, which 

suggests a possible linkage between QpZAG46 and genes encoding for leaf size.

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contrasting environments. Analyses of 3-year-old seedlings 

revealed population differentiation for eight variables: bud 

burst, bud set, multiple flushing, height, deviation from regression of 3-year height on 2-year height, spring frost damage, 

fall frost damage, and winter injury. These analyses, as well 

as high intercorrelations among population means, suggested 

that adaptations result from a balance between selection for a 

high growth potential in mild environments and selection for 

cold hardiness in severe environments. Consequently, genetic 

variation among populations was closely related to the elevation, geography, and climate of the seed source. 

Link: http://www.treesearch.fs.fed.us/pubs/38632

740. Rehfeldt GE. 1985. Ecological genetics of Pinus 

contorta in the Wasatch and Uinta Mountains of Utah. Canadian 

Journal of Forest Research 15:524-530.

Type: Journal 

Geographic Area: Utah, USA

Compilers’ Keywords: lodgepole pine, genetic differentiation, shoot elongation, genetic x environment interaction, seed 

transfer

Abstract: Genetic differentiation of 45 populations of Pinus 

contorta primarily from the Wasatch and Uinta Mountains was 

studied in field, greenhouse, and laboratory tests. Analyses of 

variables reflecting growth potential, morphology, cold hardiness, and periodicity of shoot elongation revealed population 

differentiation for a variety of traits. Regression models related 

as much as 77% of the variance among population means to 

the elevation and geographic location of the seed source. For 

genetic variation to be arranged along relatively steep environmental clines implies pronounced adaptive differentiation. 

As a result, seed transfer in reforestation should be restricted 

severely if maladaptation is to be controlled. 

741. Rehfeldt GE. 1985. Genetic variances and covariances 

in Pinus contorta: estimates of genetic gains from index selection.

Silvae Genetica 34:26-33.

Type: Journal 

Geographic Area: Pacific Northwest USA

Keywords: Pinus contorta, additive genetic variances, genetic 

correlations, genetic gains, restricted selection indices

Summary: Genetic variances and covariances were estimated 

for 6-year-old trees from 10 families within eight populations 

of Pinus contorta from the northern Rocky Mountains (USA). 

Analyses of 10 traits reflecting growth, quality, and adaptedness revealed substantial genetic variances for nearly all traits. 

As a consequence, univariate selection for height should provide genetic gains of about 5% per unit selection intensity. 

However, strong genetic correlations linked height with branch 

length (0.75), crown width (0.75), and the amount of shoot 

elongation that continues into mid-summer (0.70). Height was 

significant differences among populations in six traits: bud 

burst, bud set, 3-year height, growth rate, cold injury, and flushing periods during the first or third growing season. Most traits 

were highly intercorrelated; thus, populations that expressed 

a high growth potential also set buds late, flushed twice, grew 

at a rapid rate, but were most severely injured by freezing. 

Consequently, adaptation of populations for numerous traits is 

viewed as a balance between selection of high growth potential 

in relatively mild environments and selection for cold hardiness in severe environments. Multiple regression analyses related genetic differentiation of populations to geographic and 

ecologic conditions of the seed source. The regression model 

that best fit the data described adaptive variation according 

to two physiographic zones, elevation within each zone, and 

latitude within zones. Patterns of adaptive differentiation were 

used to develop seed transfer guidelines for reforestation and 

tree improvement.

Link: http://www.treesearch.fs.fed.us/pubs/37659

738. Rehfeldt GE. 1983.Genetic variability within Douglasfir populations: implications for tree improvement. Silvae 

Genetica 32:9-14.

Type: Journal 

Geographic Area: Idaho, Montana, USA

Keywords: Pseudotsuga menziesii, adaptive strategy, adaptation

Abstract: Genetic variances and covariances for growth potential, phenology and patterns of first year elongation were 

calculated from 30 half-sib families from each of three contrasting populations. Analyses of 4-year old trees growing in 

a single environment revealed high levels of additive genetic 

variance within populations. As a consequence, rather high 

estimates of genetic gains in growth potential were associated with weak selection intensities. However, genetic correlations were strong. Gains in growth potential were associated 

with delayed bud set and increased susceptibility to early fall 

frosts. For tree improvement to increase the growth potential 

of Douglas-fir without inadvertent degeneration of adaptation, 

selections must be based on several traits. 

Link: http://sauerlaender-verlag.com/fileadmin/content/

dokument/archiv/silvaegenetica/32_1983/32-1-2-9.pdf

739. Rehfeldt GE. 1983. Ecological adaptations in Douglasfir (Pseudotsuga menziesii var. glauca) populations. III. Central 

Idaho. Canadian Journal of Forest Research 13:626-632.

Type: Journal 

Geographic Area: Idaho, USA

Compilers’ Keywords: common garden, genetic variability, 

seed source, genetic × environment interaction, phenology

Abstract: Growth, phenology, and cold hardiness of seedlings from 74 populations of Douglas-fir (Pseudotsuga menziesii var. glauca) from central Idaho were compared in four 

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when 3-year height was expressed as the growth from a common height at age 2. Greenhouse studies of the periodicity of 

shoot elongation revealed that populations of high growth potential achieved a large stature by means of a long duration, late 

cessation, rapid rate, and large amount of elongation. Multiple 

regression models accounted for as much as 60 percent of the 

variance between populations and described adaptive landscapes in which populations from low elevation have a high 

growth potential while those from high elevations have a low 

growth potential. Geographic variation at a common elevation indicated that populations of high growth potential come 

from areas where the frostfree period is long and precipitation 

is relatively high. Thus, growth potential is directly related to 

the length of the growing season, which can be truncated by 

either frost or moisture stress. Adaptive landscapes are used 

to construct guidelines for limiting seed transfer in artificial 

reforestation. In general, seed from a single source should not 

be transferred more than ±200 m in elevation.

Link: http://www.treesearch.fs.fed.us/pubs/38597

744. Rehfeldt GE. 1988. Ecological adaptations in Douglasfir (Pseudotsuga menziesii var. glauca). IV. Montana and Idaho 

near the Continental Divide. Western Journal of Applied Forestry 

3:101-105.

Type: Journal 

Geographic Area: Idaho, Montana, USA

Compilers’ Keywords: genetic differentiation, growing season length, genetic x environment interaction, cline, seed transfer guidelines

Abstract: Seventy-seven seedlings populations of Douglasfir (Pseudotsuga menziesii var. glauca) sampled from near the 

Continental Divide in Idaho and Montana exhibited pronounced 

genetic differences when compared in three common environments. Differentiation involved several traits several traits 

that are components of an annual developmental cycle that 

must be completed within a growing season of finite length. 

Consequently, elevational and geographic clines of genetic 

variation tend to parallel variation in the length of the growing 

season at the seed source. Such clines suggest that maladaptation in artificial reforestation can be controlled by limiting the 

transfer of seeds. While geographic transfers can be relatively 

liberal, elevational transfers should be limited to ±90 m (300 ft) 

from the seed source at elevations below 1,400 m (4,600 ft), 

to ±125 m (410 ft) for elevations between 1,400 and 2,000 m 

(4,600 and 6,550 ft), and ±200 m (560 ft) for elevations above 

2,000 m (6,550 ft).

Link: http://www.treesearch.fs.fed.us/pubs/38630

745. Rehfeldt GE. 1988. Ecological genetics of Pinus 

contorta from the Rocky Mountains (USA): a synthesis. Silvae 

Genetica 37:131-135.

Type: Journal 

also related to the number of whorls on the current shoot and, 

consequently, to the number of branches. Thus, multi-trait selections are necessary to prevent genetic gains in productivity 

from being accompanied by inadvertent degeneration of traits 

related to adaptation and quality. Multi-trait selections, however, reduce anticipated genetic gains in productivity. Restricted 

selection indices are developed to exemplify procedures for 

holding constant the correlated responses while maximizing 

gains in height. 

Link: http://silvaegenetica.com/fileadmin/content/dokument/archiv/silvaegenetica/34_1985/34-1-26.pdf

742. Rehfeldt GE. 1986. Adaptive variation in Pinus ponderosa from Intermountain regions. I. Snake and Salmon River 

Basins. Forest Science 32:79-92.

Type: Journal 

Geographic Area: Idaho, USA

Keywords: genetic variation, population differentiation, ecological genetics, genecology

Abstract: Genetic differentiation of 64 populations from central Idaho was studied in field, greenhouse and laboratory tests. 

Analyses of variables reflecting growth potential, phenology, 

morphology, cold hardiness and periodicity of shoot elongation revealed population differentiation for a variety of traits. 

Regression models related as much as 61 percent of the variance among populations to the elevation and geographic location of the seed source. Clinal patterns of adaptive variation 

provide the basis for developing seed transfer guidelines that 

will control maladaptation in reforestation. In central Idaho, 

for example, seed transfer should be limited to within 180 m 

of the seed source.

Link: http://www.treesearch.fs.fed.us/pubs/38631

743. Rehfeldt GE. 1986. Adaptive variation in Pinus ponderosa from intermountain regions. II. Middle Columbia River 

System. Ogden (UT): USDA Forest Service, Intermountain 

Research Station. RP-INT-373. 12 p.

Type: Government Document

Geographic Area: Northwestern USA

Compilers’ Keywords: ponderosa pine, adaptive variation, 

drought tolerance, common garden, seed transfer guidelines, 

growing season length

Research Summary: Seedlings representing 138 populations 

of ponderosa pine from drainages of the middle Columbia 

River system were grown and compared in common environments. Field studies of 3-year-old trees on mesic and xeric test 

sites detected population differentiation for traits reflecting 

growth and development. Populations that expressed the greatest growth potential under mesic culture suffered the greatest 

reduction in height under xeric culture. Consequently, a different set of populations was performing the best on the two sites 
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Geographic Area: Southwestern USA

Compilers’ Keywords: ponderosa pine, common garden, 

genetic variability, genetic × environment interaction, clinal variation

Abstract: Genetic variation among 62 populations of 

ponderosa pine was studied by comparing seedlings from 

all populations according to (1) growth and development 

of 4-yr-old seedlings in three disparate common gardens 

and (2) patterns of shoot elongation of 2-yr-old seedlings 

in a greenhouse. Genetic variation was detected among 

populations for 19 of the variables, most of which were 

intercorrelated. Two principal components accounted for 

60% of the total interpopulation variance. Multiple regression analyses were used to relate genetic variation in 19 

variables and two principal components to the elevation 

and geographic origin of the seed. The regression models 

produced values of R2 as large as 0.78 and accounted for 

more than 40% of the variance among populations for 14 

of the variables, including both the first and second principal components. These models described genetic variation 

as occurring along a relatively steep elevational cline and 

along both primary and secondary geographic clines of 

relatively gentle slope. All clines paralleled patterns of environmental variation, particularly the length of the frostfree season and patterns of precipitation. Because genetic 

variation occurs along three clines simultaneously, genetic 

differentiation can be described as rampant. Nevertheless, 

similar genotypes tend to recur in similar environments.

Link: http://www.treesearch.fs.fed.us/pubs/38628

748. Rehfeldt GE. 1991. A model of genetic variation for 

Pinus ponderosa in the Inland Northwest (USA): applications in 

gene resource management. Canadian Journal of Forest Research 

21:1491-1500.

Type: Journal 

Geographic Area: Northwestern USA

Compilers’ Keywords: ponderosa pine, common garden, genetic x environment interaction, gene conservation, seed zones, 

seed transfer, phenotypic variation, breeding zones

Abstract: Models were developed to describe the genetic 

variation among 201 seedling populations of Pinus ponderosa

var. ponderosa in the Inland Northwest of the United States. 

Common-garden studies provided three variables that reflected 

growth and development in the field environments and three 

principal components of six variables that reflected patterns 

of shoot elongation. Regression models were developed for 

describing genetic variation across the landscape. Using functions of latitude, longitude, and elevation as descriptors, these 

models produced values of R2 that were as large as 0.66, while 

averaging 0.39. The models described genetic variation as occurring along relatively steep elevational clines and gentle geographic (i.e., latitudinal and longitudinal) clines. An exercise 

Geographic Area: Rocky Mountains USA

Keywords: population differentiation, genecology, adaptive 

variation

Abstract: Studies of population differentiation in Pinus contorta, which were conducted in common gardens, are summarized according to the growth, freezing tolerance, and the 

periodicity of shoot elongation of seedlings from 173 Rocky 

Mountain (USA) populations. Regression models accounted 

for 45 to 77% of the variance among populations, coordinated 

the results of previous studies and described clinal patterns of 

variation that generally reflect elevational and geographic gradients in the frost-free period. 

Link: http://germanjournalofforestresearch.com/

fileadmin/content/dokument/archiv/silvaegenetica/37_1988/37-3-4-131.pdf

746. Rehfeldt GE. 1989. Ecological adaptations in Douglasfir (Pseudotsuga menziesii var. glauca): a synthesis. Forest Ecology 

and Management 28:203-215.

Type: Journal 

Geographic Area: Pacific Northwest USA

Compilers’ Keywords: common garden, genetic variation, 

seed transfer

Abstract: Measurements of 3rd-year height of 228 seedling 

populations, grown in four separate studies in two of the same 

common gardens, were used to summarize patterns of genetic 

variation for Douglas-fir across 250,000 km2 of forested lands 

in Idaho and Montana, U.S.A. Because each study was conducted in different years with a different set of populations, 

measurements were transformed to standard deviates and then 

were scaled according to the performance of populations common between studies. Genetic variation in 3rd-year height was 

related to the elevation and geographic location of the seed 

source by a regression model that accounted for 87% of the 

variance among populations. In addition, 3rd-year height of 

169 of the populations was strongly correlated (r = 0.80) to 

freezing injury observed in previous studies. Both variables 

showed that populations from elevational or geographically 

mild sites were tall but had low freezing tolerance. Populations 

from harsh sites were short and cold hardy. In Douglas-fir, 

adaptation to heterogeneous environments can be viewed as 

physiological specialization for a relatively small portion of 

the environmental gradient; populations separated by a relatively short distance along the environmental gradient (e.g., 20 

frost-free days) tend to be different genetically.

Link: http://www.treesearch.fs.fed.us/pubs/38629

747. Rehfeldt GE. 1990. Genetic differentiation among 

populations of Pinus ponderosa from the Upper Colorado River 

Basin. Botanical Gazette 151:125-137.

Type: Journal 
264 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

500 m in elevation before genetic differentiation can be expected. Regression models using geographic predictors were 

developed to describe variation in ten climatic variables from 

192 weather stations that best represented the Rocky Mountain 

distribution of L. occidentalis. Values of R2 ranged from 0.43 

to 0.92 and were higher for temperature than for precipitation 

variables. Using climatic effects predicted for each provenance 

to describe genetic variation produced values of R2 between 

0.15 and 0.27 which accounted for nearly as much (68-100%) 

of the genetic variance as had geographic variables. The analyses suggested that genetic differentiation was controlled primarily by the relative mildness of the climate. Secondary effects of precipitation were implicated for variables measuring 

the impact of Meria needle cast and survival.

751. Rehfeldt GE. 1999. Systematics and genetic structure of Washoe pine: applications in conservation genetics.

Silvae Genetica 48:167-173.

Type: Journal 

Geographic Area: Western North America

Keywords: Pinus washoensis, quantitative traits, genetic structure, systematics, conservation biology

Abstract: Independent studies of seedling populations of 

Washoe (Pinus washoensis) and ponderosa (P. ponderosa) 

pines grown in common gardens demonstrated that: (1) the 

systematic relationship between Washoe pine and the North 

Plateau race of ponderosa pine is close and (2) the allocation 

of genetic variability among and within populations of the narrow endemic, Washoe pine, is similar to that of the broadly 

dispersed ponderosa pine. The results from this quantitative 

analysis of adaptive traits thus support previous works involving morphology, terpene chemistry, allozyme variation, mating 

systems, DNA biochemistry, and classical taxonomy that lead 

to a conclusion of synonymity for Washoe pine and ponderosa 

pine. The results also provide no genetic evidence that small 

population sizes and isolated distributions have had deleterious 

genetic consequences. Populations of Washoe pine nevertheless have unique characteristics that may be worthy of conservation. Programs should concentrate on habitat preservation 

and range expansion.

Link: http://silvaegenetica.com/fileadmin/content/dokument/archiv/silvaegenetica/48_1999/48-3-4-167.pdf

752. Rehfeldt GE. 2001. Introduction of ponderosa pine 

and Douglas-fir to Argentina. New Forests 21:35-44.

Type: Journal 

Geographic Area: USA, Argentina

Keywords: afforestation, land races, provenance selection, reforestation, shoot elongation, tree introduction

Abstract: Patterns of shoot elongation of 2-yr seedlings from 

native North American populations of ponderosa pine and 

at validating the models with independent data supported their 

veracity. Predictions made by the models are applied to limiting seed transfer, designing breeding zones, planning gene 

conservation programs, interpreting phenotypic variation, and 

predicting the effects of environmental change on the adaptedness of populations. 

Link: http://www.treesearch.fs.fed.us/pubs/38627

749. Rehfeldt GE. 1993.Genetic variation in the Ponderosae 

of the Southwest. American Journal of Botany 80:330-343.

Type: Journal 

Geographic Area: Southwestern USA

Compilers’ Keywords: Pinus ponderosa, common garden, 

genetic × environment interaction, population differentiation, 

model application, seed transfer guidelines, seed zones

Abstract: Ninety-five seedling populations of southwestern 

ponderosa pine (Pinus ponderosa var. scopulorum) along with 

single populations of Pinus engelmannii and Pinus arizonica

were compared in four environmentally disparate common 

gardens. Differentiation among ponderosa pine populations 

was detected for a diverse assortment of variables that included patterns of shoot elongation, measure of growth potential, 

winter and spring freezing damage, and leaf characteristics. 

Multiple regression models accounted for as much as 85% 

of the variance among populations and described complex 

clines that were dominated by elevational and latitudinal effects. Although P. ponderosa, P. arizonica, and P. engelmannii

were readily differentiated, the performance of progenies from 

one population suggested introgression primarily involving P. 

ponderosa and P. arizonica but also implicating P. engelmannii. 

Link: http://www.treesearch.fs.fed.us/pubs/38626

750. Rehfeldt GE. 1995. Genetic variation, climate models 

and the ecological genetics of Larix occidentalis. Forest Ecology 

and Management 78:21-37.

Type: Journal 

Geographic Area: Northwestern North America

Keywords: population differentiation, climate patterns, 

genecology

Abstract: Provenance tests of 143 populations of Larix 

occidentalis revealed genetic differentiation for eight variables 

describing growth, phenology, tolerance to spring frosts, effects of Meria laricis needle cast, and survival. Geographic 

variables accounted for as much as 34% of the variance among 

Rocky Mountain populations. Patterns of genetic variation 

were dominated by the effects of latitude and elevation, with 

populations from the north and from high elevations having 

the lowest growth potential, the least tolerance to the needle 

cast, and the lowest survival. However, the slope of the geographic clines was relatively flat. Populations in the same 

geographic area, for instance, need to be separated by about 

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 265

Geographic Area: Western USA

Compilers’ Keywords: western white pine, geographic pattern, genetic variability, common garden, genetic × environment 

interaction

Abstract: Genetic differentiation of 59 populations, representing the geographic distribution of Pinus monticola, was 

studied in field, greenhouse, and laboratory tests. Geographic 

variation was described by regression models, which accounted for as much as 85% of the variance among populations. Populations from the north (Rocky Mountains, northern 

Cascades, and northern coastal areas) are generally of high 

growth potential and low cold hardiness. Southern populations 

(Sierra Nevada) exhibit low growth potential and high hardiness. Populations from the central and southern Cascades are 

arranged along relatively steep latitudinal lines that connect 

northern and southern groups. Although differentiation within 

the transitional region was readily detected, patterns of variation within northern and southern regions were either weak 

or nonexistent. Nowhere was genetic variation related to the 

elevation of the seed source.

755. Rehfeldt GE, Tchebakova NM, Milyutin LI, 

Parfenova EI, Wykoff WR, Kouzmina NA. 2003.

Assessing populations responses to climate in Pinus sylvestris 

and Larix spp. of Eurasia with climate-transfer models. Eurasian 

Journal of Forest Research 6:83-98.

Type: Journal 

Geographic Area: USA, Canada, Russia

Keywords: climate change responses, climatypes, genetic variation, reforestation, seed transfer guidelines

Abstract: Weibull regression models were used to relate 

height and survival of Eurasian populations of Scots pine 

(Pinus sylvestris L.) at age 13 and three species of larch (L. 

sukaczewii Dylis, L. sibirica Ledeb., and L. gmelinii (Rupr.) 

Rupr.) at age 12 to the difference in climate between their 

provenance and a planting site. Univariate models using 

five climate variables as predictors all were statistically significant (p < 0.01), and all but the pine survival functions 

received strong verification with independent data. The 

models showed that the growth and survival of most populations of the pine and each species of larch are enhanced 

when populations are transferred from their provenance to 

warmer climates. The results are consistent with the view 

that most populations occur in climates that are suboptimal, 

the degree of which is directly related to the severity of the 

climate. Because of this, projected responses to a climatechange scenario of the Hadley Centre were highly variable 

geographically. Short-term plastic responses tended to be 

strongly negative for the least severe climates and strongly 

positive for the most severe. Long-term evolutionary responses primarily reflected extirpation and immigration for 

the species of larch but showed additionally for the pine that 

Douglas-fir were compared to those of Argentine land races 

originating from unknown provenances. The comparisons 

were conducted in Moscow, Idaho (USA), and suggested that 

the ponderosa pine land race was descended from a California 

provenance at low or middle elevations but that the growth 

potential of the land race was only mediocre in comparison to 

eight native populations. The Douglas-fir land race exhibited 

a relatively high growth potential in comparison to 19 native 

provenances and undoubtedly originated from a mild coastal 

environment. The results provide concrete recommendations 

for upgrading the growth potential of the Argentine land races 

by importing germplasm of specific provenances.

Link: http://www.treesearch.fs.fed.us/pubs/38625

753. Rehfeldt GE. 2005. Geographic clines in genetic variation. In: Mardsen M, Downing M, Riffe M, editors. Workshop 

Proceedings: Quantitative Techniques for Deriving National 

Scale Data. Fort Collins (CO): USDA Forest Service, Forest 

Health Technology Enterprise Team. FHTET-2005-12. p 

223-261.

Type: Government Document

Geographic Area: Western North America

Compilers’ Keywords: Pseudotsuga menziesii, genetic variation, geographic clines, risk mapping, climate modeling, seed 

zone delineation, seed transfer guidelines, climate change

Abstract: In risk mapping, the primary considerations are the 

presence of the host tree species, some measure of its density, 

and the distribution of the pest agent. High-density or overstocked stands are often considered to be of higher risk then 

stands with lower stocking levels; also important is the climatic stress on the population. This presentation shows how the 

predictions from a climate model can be converted to variables 

that may indicate the status of the stress of conifer species 

and their populations in the western USA and southwestern 

Canada. Forty-eight monthlies were derived from the basic 

temperature and precipitation data and fit to geographic surfaces with thin plate splines. These monthlies were then used 

to describe the clines of genetic variation that exist within species for growth characteristics. The mapping of clinal variation 

is useful in delineating seed zones and deriving seed transfer 

guidelines. The reverse image of such maps should indicate 

where the species would be under stress due to climatic conditions. For this reason, it is recommended that the monthlies and 

the climatic limits could be useful in risk mapping.

Link: http://www.fs.fed.us/foresthealth/technology/pdfs/

QuantitativeTechniques.pdf

754. Rehfeldt GE, Hoff RJ, Steinhoff RJ. 1984.

Geographic patterns of genetic variation in Pinus monticola.

Botanical Gazette 145:229-239.

Type: Journal 

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266 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

to climate similarities of the provenances to provide single 

datasets for 781 P. ponderosa and 1193 P. menziesii populations. Mixed effects models were used for two sub-specific 

varieties of each species to describe clines in growth potential associated with provenance climate while accounting for 

study effects not eliminated by scaling. Variables related to 

winter temperatures controlled genetic variation within the 

varieties of both species. Clines were converted to climatypes by classifying genetic variation, using variation within 

provenances in relation to the slope of the cline to determine 

climatype breadth. Climatypes were broader in varieties of P. 

ponderosa than in P. menziesii and were broader for varieties 

inhabiting coastal regions of both species than for varieties 

from interior regions. Projected impacts of climate change 

on adaptedness used output from an ensemble of 17 general 

circulation models. Impacts were dependent on cline steepness and climatype breadth but implied that maintaining 

adaptedness of populations to future climates will require a 

redistribution of genotypes across forested landscapes.

Link: http://www.treesearch.fs.fed.us/pubs/46901

758. Rehfeldt GE, Jaquish BC, Sáenz-Romero C, 

Joyce DG, Leites LP, Bradley St. Clair J, López-Upton 

J. 2014. Comparative genetic responses to climate in the varieties of Pinus ponderosa and Pseudotsuga menziesii: reforestation.

Forest Ecology and Management 324:147-157.

Type: Journal 

Geographic Area: Western North America

Keywords: management strategies, climate-change impacts, 

Douglas-fir, ponderosa pine, seed transfer guidelines

Abstract: Impacts of climate change on the climatic niche of 

the sub-specific varieties of Pinus ponderosa and Pseudotsuga 

menziesii and on the adaptedness of their populations are 

considered from the viewpoint of reforestation. In using climate projections from an ensemble of 17 general circulation 

models targeting the decade surrounding 2060, our analyses 

suggest that a portion of the lands occupied today primarily by coastal varieties of each species contain genotypes 

that should remain suitable for the future climate. A much 

larger portion, particularly for varieties occupying inland 

sites, should require either introduction of better suited species or conversion to better adapted genotypes. Regeneration 

strategies are considered with the goal of matching growth 

potential of contemporary populations to the future climate 

where that potential can be realized. For some lands, natural 

reproduction should be suitable, but most lands will require 

forest renewal to maintain forest health, growth, and productivity. Projected impacts also illustrate the urgent need for 

conservation programs for P. menziesii in Mexico.

Link: http://www.treesearch.fs.fed.us/pubs/46902

the accommodation of global warming will require a redistribution of genotypes throughout the species’ range.

Link: http://133.87.26.249/dspace/bitstream/2115/22164/1/6(2)_P83-98.pdf

756. Rehfeldt GE, Jaquish BC, López-Upton J, 

Sáenz-Romero C, St. Clair JB, Leites LP, Joyce DG. 

2014. Comparative genetic responses to climate for the varieties 

of Pinus ponderosa and Pseudotsuga menziesii: realized climate 

niches. Forest Ecology and Management 324:126-137.

Type: Journal 

Geographic Area: Western North America

Keywords: niche modeling, climate change impacts, Douglasfir, ponderosa pine, biogeography, genecology

Abstract: The Random Forests classification algorithm was 

used to predict the occurrence of the realized climate niche 

for two sub specific varieties of Pinus ponderosa and three 

varieties of Pseudotsuga menziesii from presence-absence 

data in forest inventory ground plots. Analyses were based 

on ca. 271,000 observations for P. ponderosa and ca. 426,000 

observations for P. menziesii, with ca. 6% of the observations 

in each dataset recording the presence of one of the varieties. 

Classification errors to the respective databases attributable 

to fitting the models were ca. 5%, most of which were from 

falsely predicting varietal occurrence. Confusion in classifying varieties was nil. The primary drivers of the niche model 

were summer precipitation, winter precipitation and summer 

degree-days >5  ̊C for the varieties of P. ponderosa and the 

summer-winter temperature differential, summer maximum 

temperatures and summer precipitation for the varieties of P. 

menziesii. Projected impacts of global warming using output from an ensemble of 17 general circulation models were 

greater for P. ponderosa than for P. menziesii and for varieties of both species from inland climates than from coastal. 

Projected impacts imply dire consequences for the varieties 

of P. menziesii occurring in Mexico.

Link: http://www.treesearch.fs.fed.us/pubs/46900

757. Rehfeldt GE, Leites LP, Bradley St. Clair J, 

Jaquish BC, Sáenz-Romero C, López-Upton J, Joyce 

DG. 2014. Comparative genetic responses to climate in the varieties of Pinus ponderosa and Pseudotsuga menziesii: clines in 

growth potential. Forest Ecology and Management 324:138-146.

Type: Journal 

Geographic Area: Western North America

Keywords: genetic variation, climate-change impacts, genecology, mixed effects models, provenance tests

Abstract: Height growth data were assembled from 10 

Pinus ponderosa and 17 Pseudotsuga menziesii provenance 

tests. Data from the disparate studies were scaled according 

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Type: Journal 

Geographic Area: Western North America

Keywords: Chamaecyparis nootkatensis, yellow cedar, genetic 

structure, inbreeding, phylogeography, isozymes

Abstract: A range-wide survey of isozyme variation in yellow 

cedar (Chamaecyparis nootkatensis (D. Don) Spach; or Alaska 

cedar) found significant inbreeding and a striking geographical 

structure. The proportion of diversity among populations (GST) 

was 0.139, and three major geographical groups were evident: 

(1) Vancouver Island – mid-north coast British Columbia 

(B.C.); (2) south coast B.C. – Washington state; (3) southeast 

Alaska. This may indicate the presence of multiple refugia during ice-age range contractions. Geographically disjunct populations in Oregon and southeastern B.C. were also genetically 

distinct. Adult inbreeding coefficients (F) were usually positive 

but varied widely among populations, as did gene diversity. 

The overall average of F = 0.18 suggests that considerable selfing (ca. 30%) occurs in this species. Other statistics were more 

typical for a conifer: within population heterozygosity (HS) 

averaged 0.148, total gene diversity (HT) was 0.171, and the 

number of alleles per locus and percentage polymorphic loci 

averaged 1.68 and 50, respectively. This geographic structure 

has implications for species conservation and for breeding programs. Also, previous studies of other Cupressoideae (Rich. 

ex Sweet) species indicate that these levels of inbreeding and 

population structure are not uncommon, and may influence 

evolutionary divergence in the Cupressoideae.

Link: http://www.genetics.forestry.ubc.ca/ritland/reprints/2001_CJB_YellowCedarIsozymes.pdf

761. Roberds JH, Conkle MT. 1984. Genetic structure 

in loblolly pine stands: allozyme variation in parents and progeny.

Forest Science 30:319-329.

Type: Journal 

Geographic Area: Southeastern USA

Keywords: population subdivision, allele frequencies, F statistics, isozymes, Pinus taeda

Abstract: Genetic population structure in two adjacent 

old-field stands of loblolly pine was studied by analyzing 

allele frequencies for eight polymorphic isozyme loci. The 

stands represented different generations with trees in the 

older stand being parents of trees in the younger one. The 

parental stand consisted of three fairly distinct age classes 

which differed significantly in allele frequencies for four of 

the allozyme loci, yet no differences were found among the 

spatial divisions studied. Within the progeny stand, no genetic differentiation was detected with respect to either age 

class or spatial grouping. Although allele frequencies did not 

differ between the two stands, genetic population structure 

was not the same, demonstrating that local genetic structure 

can differ between successive generations and that varying 

759. Richardson BA, Kitchen SG, Pendleton RL, 

Pendleton BK, Germino MJ, Rehfeldt GE, Meyer 

SE. 2014. Adaptive responses reveal contemporary and 

future ecotypes in a desert shrub. Ecological Applications 

24:413-427.

Type: Journal 

Geographic Area: Southwestern USA

Keywords: assisted migration, blackbrush, climate change, 

Coleogyne ramosissima, ecological restoration, seed transfer 

zones

Abstract: Interacting threats to ecosystem function, including climate change, wildfire, and invasive species necessitate 

native plant restoration in desert ecosystems. However, native 

plant restoration efforts often remain unguided by ecological genetic information. Given that many ecosystems are in 

flux from climate change, restoration plans need to account 

for both contemporary and future climates when choosing 

seed sources. In this study we analyze vegetative responses, 

including mortality, growth, and carbon isotope ratios in two 

blackbrush (Coleogyne ramosissima) common gardens that 

included 26 populations from a range-wide collection. This 

shrub occupies ecotones between the warm and cold deserts 

of Mojave and Colorado Plateau ecoregions in western North 

America. The variation observed in the vegetative responses 

of blackbrush populations was principally explained by grouping populations by ecoregions and by regression with site-specific climate variables. Aridity weighted by winter minimum 

temperatures best explained vegetative responses; Colorado 

Plateau sites were usually colder and drier than Mojave sites. 

The relationship between climate and vegetative response was 

mapped within the boundaries of the species-climate space 

projected for the contemporary climate and for the decade surrounding 2060. The mapped ecological genetic pattern showed 

that genetic variation could be classified into cool-adapted and 

warm-adapted ecotypes, with populations often separated by 

steep clines. These transitions are predicted to occur in both the 

Mojave Desert and Colorado Plateau ecoregions. While under 

contemporary conditions the warm-adapted ecotype occupies 

the majority of climate space, climate projections predict that 

the cool-adapted ecotype could prevail as the dominant ecotype as the climate space of blackbrush expands into higher 

elevations and latitudes. This study provides the framework 

for delineating climate change-responsive seed transfer guidelines, which are needed to inform restoration and management 

planning. We propose four transfer zones in blackbrush that 

correspond to areas currently dominated by cool-adapted and 

warm-adapted ecotypes in each of the two ecoregions.

Link: http://www.treesearch.fs.fed.us/pubs/45555

760. Ritland C, Pape T, Ritland K. 2001. Genetic structure of yellow cedar (Chamaecyparis nootkatensis). Canadian 

Journal of Botany 79:822-828.

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purposes. Individual populations across different valleys 

also proved to be a valuable source of variation for economically useful traits. A discussion on possible implications of 

use of nonlocal germplasm sources for restoration purposes 

is provided. The lowland site gave much lower seed production than the mountain location and experienced outstanding plant mortality after the first summer. The specific site 

conditions, namely the occurrence of an appreciable level of 

heat and drought stress in summer and a severe rust infection 

rather than the altitude per se, limited the suitability of this 

site for seed production of the alpine germplasm.

764. Roy BA, Mulder CPH. 2014. Pathogens, herbivores, and phenotypic plasticity of boreal Vaccinium vitis-idaea

experiencing climate change. Ecosphere 5:1-19.

Type: Journal 

Geographic Area: Alaska, USA

Keywords: Alaska, climate change, Exobasidium vaccinia, 

global warming, herbivory, local adaptation, lingonberry, 

Phenolic, Phomopsis columnaris, plasticity, reciprocal transplant, 

Vaccinium vitis-idaea

Abstract: Climate warming is occurring at a rapid rate in 

the boreal forest; mean winter temperature has increased 

about 4 °C in Alaska over the last 40 years and about the 

same increase is predicted over the next 40 years. Warming 

temperatures tend to increase the number and kinds of herbivores and pathogens. How will boreal plants to respond 

to these abiotic and biotic changes? To address these questions we used common gardens and reciprocal transplants of 

Vaccinium vitis-idaea at sites with contrasting abiotic conditions near Fairbanks, Alaska. Plant morphology, chemistry, 

resistance to pathogens and herbivores, and survival were all 

strongly influenced by the destination environment (planting 

site and block within site), and less by the site of origin for 

the seed. Overall, seedlings survived significantly better at 

the summer cold site, which was buffered from drought as 

a result of its northern aspect, presence of sphagnum moss 

and permafrost (86.4% survival versus 48.5%). However, 

this cool damp site had more stem-killing consumers, the 

pathogens Phomopsis columnaris and Exobasidium vaccinii

and mammalian browse, all of which became more common 

over the three years of the study as the plants became larger. 

Taken together, these results suggest that the seedling stage 

is likely to be vulnerable to droughts, except at sites with 

thick moss cover, which results in greater duration of summer moisture. Thus, as the climate warms, seedling growth 

will be restricted to those sites where the adult stages will 

later suffer from more stem-killing pathogen attack and 

mammalian browse.

Link: http://www.esajournals.org/doi/abs/10.1890/ES13-

00271.1

structures can exist in localized regions. Evidence suggesting a low level of inbreeding was found in both stands.

762. Rogers DL, Harry DE, Libby WJ. 1994. Genetic 

variation in incense-cedar (Calocedrus decurrens): I. provenance 

differences in a twelve-year-old common-garden study. Western 

Journal of Applied Forestry 9:113-117.

Type: Journal 

Geographic Area: Oregon, California, USA

Compilers’ Keywords: tree improvement, provenance test

Abstract: Results are reported from a provenance study of 

incense-cedar (Calocedrus decurrens), a tree species native to 

forested regions of Oregon, California, and Baja California. 

Seedlings from six geographically distinct regions were 

measured over a 12 yr period in a common-garden study located near the center of the species’ native range. In general, 

among-region genetic variation was small. However, differences in height, stem volume, and crown form between trees 

from the southernmost region sampled and trees from the 

other five study regions were large and significant. Further 

studies of within-population genetic variation are recommended prior to initiating any tree improvement activities. 

763. Romani M, Pecetti L, Tosca A, Della 

Marianna G, Gusmeroli F, Paoletti R, Piano E. 

2009. Evaluation of morphology and seed production of alpine 

cat’s tail [Phleum rhaeticum (Humph.) Rauschert] germplasm 

in view of its use for ecological restoration at high altitude.

Restoration Ecology 17:386-392.

Type: Journal 

Geographic Area: Italy, Europe

Keywords: genetic resources, high-altitude restoration, Phleum 

rhaeticum, seed production, selection, ‘‘site-specific’’ vegetation

Abstract: ‘‘Site-specific’’ vegetation, that is germplasm 

ecologically adapted to the prevailing pedoclimatic conditions and native to the target region, is increasingly required 

in the revegetation of disturbed areas at high altitude. Some 

key site-specific species have been reported for the Alpine 

region. Better knowledge is needed on the existing variability of these species, while high seed yield is required to ensure the availability of commercially adequate seed supply. 

Natural populations of Alpine cat’s tail from the Rhaetian 

Alps were evaluated for morphology and seed production at 

a mountain and at a lowland site of northern Italy aiming at 

(1) identifying possible superior populations for the selection 

of adapted high yielding cultivars and (2) verifying whether 

the seed multiplication of site-specific species could also be 

undertaken in areas different from those of origin. The germplasm from one valley featured an interesting architecture of 

traits, representing a possible ideotype for seed production 

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sites located south of latitude 56° N; a high-elevation (1,220 

m) site was poorly correlated (rp ≤ 0.40) with most of the sites 

located at medium and low elevations (≤ 800 m); additive 

main effects and multiplicative interaction analysis revealed a 

strong association between provenances and sites with similar 

climate; high-latitude provenances and sites with cool winters 

and dry summers, and high elevation provenances and sites 

with high precipitation and short growing seasons contributed 

the greatest to GEI. Because the relationship between GEI and 

climate corresponds well with latitudes and topography, seed 

transfer in Alberta can be effectively regulated by setting latitude and elevation transfer limits that are periodically adjusted 

to reflect the changing conditions caused by climate change.

767. Rweyongeza DM, Barnhardt LK, Hansen CR. 

2011. Patterns of optimal growth for white spruce provenances in Alberta. Smoky Lake, Alberta, Canada: Government 

of Alberta, Sustainable Resource Development, Alberta Tree 

Improvement and Seed Centre. Ref. T/255. 37 p.

Type: Government Document

Geographic Area: Western Canada

Compilers’ Keywords: genetic variation, climatic variability, 

seed source, seed transfer guidelines

Summary: This paper describes the pattern of genetic variation for growth potential among white spruce provenances in 

Alberta, Canada. It uses total height (H27) and diameter at 

breast height (D27) at 27-yrs from seed to identify provenance 

differences in relation to latitude and elevation of seed origin. 

Latitude and elevation are the main variables that adequately 

describe climatic variability in Alberta. Indirectly, latitude also 

describes regional variation in day length. Climate and day 

length are the main natural selection pressures governing genetic differentiation among forest tree populations. The regressions of height and diameter on latitude and elevation of seed 

origin are used to predict the location from which provenances 

of the highest growth potential on a specified site or environment are expected to originate. This location is referred to as an 

optimum seed source. By examining the pattern of provenance 

variation on similar sites and the correlation of provenance 

ranks among sites (genotype by environment interaction), it is 

possible to establish seed transfer guidelines that will enhance 

efficient use of seed while maintaining provenance climatic 

adaptation.

Link: http://srd.alberta.ca/MapsPhotosPublications/

Publications/documents/PatternsOfOptimalGrowthForWhiteSpruce-Nov2011.pdf

768. Sáenz-Romero C, Guzmán-Reyna RR, Rehfeldt 

GE. 2006. Altitudinal genetic variation among Pinus oocarpa

populations in Michoacán, Mexico: implications for seed zoning, conservation, tree breeding and global warming. Forest 

Ecology and Management 229:340-350.

765. Rukavina H, Hughes H, Johnson R. 2008.

Variation in saltgrass growth and time of fall dormancy related 

to geographical and climatic factors. Journal of the American 

Society for Horticultural Science 133:127-132.

Type: Journal 

Geographic Area: Western USA

Keywords: turfgrass breeding, phenotypic variability, cold hardiness, warm-season turfgrass

Abstract: Development of a new turfgrass cultivar requires 

an evaluation of numerous traits as well as an understanding of 

environmental factors influencing those traits. Growth or ability 

to fill in gaps and time of fall dormancy (fall color retention) 

that indicates cold hardiness are important traits for turfgrasses. 

This study was initiated to characterize variation in saltgrass 

[Distichlis spicata L. (Greene)] growth and time of fall dormancy 

related to climatic and geographical factors at the source location 

(geographical location of clone origin). Growth traits and time 

of fall dormancy were measured on 52 saltgrass clones collected 

from 41 locations and established at one location (common 

garden) in Fort Collins, CO. Principal component analysis on 

the morphological traits extracted the first principal component 

that explained 78% of the variability. The first principal component and time of fall dormancy were related to climatic and 

geographical factors at the source locations. Variation in growth 

traits was related to seasonal climatic variables of summer drying and fall cooling that explained 50% of variability in morphological traits. Variation in time of fall dormancy was related 

to longitude of clone origin and minimum winter temperature. 

These two variables explained 60% of the total variability in 

time of fall dormancy. Information obtained in this study may 

help breeders identify the best environments for specific traits 

and suggests that cold tolerance could be a problem for some 

clones from western sources if established too far east.

Link: http://www.treesearch.fs.fed.us/pubs/29634

766. Rweyongeza DM. 2011. Pattern of genotype–

environment interaction in Picea glauca (Moench) Voss in 

Alberta, Canada. Annals of Forest Science 68:245-253.

Type: Journal 

Geographic Area: Western Canada

Keywords: AMMI, climate, provenance trial, type B correlation, white spruce

Abstract: Genotype-environment interaction (GEI) among 

19 white spruce provenances planted on eight sites in Alberta, 

Canada, was analyzed using type B correlations (rp) and the 

additive main effects and multiplicative interaction (AMMI) 

model. The objectives were to quantify the extent of crossover 

interaction for height and diameter; evaluate the age trends in 

GEI for height and diameter; and examine the role of provenance and test site climate in causing GEI. A high latitude 

(~59° N) site was poorly correlated (rp = −0.31 to 0.56) with 

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769. Salmela MJ. 2011. Adaptive genetic variation in Scots 

pine (Pinus sylvestris L.) in Scotland [Dissertation]. Edinburgh, 

Scotland, United Kingdom: The University of Edinburgh. 170 p.

Type: Dissertation

Geographic Area: Scotland, United Kingdom

Compilers’ Keywords: phenotypic traits, common garden, 

genetic × environment interaction, seed transfer guidelines

Abstract: Genetic differentiation in phenotypic traits 

among populations from heterogeneous environments is 

often observed in common-garden studies on forest trees, 

but data on adaptive variation in Scots pine (Pinus sylvestris

L.) in Scotland are limited. As a result, current seed transfer 

guidelines are based on earlier molecular marker studies and 

do not take into account environmental or adaptive genetic 

variation. An analysis of spatial variation in climate showed 

substantial differences in temperature and precipitation 

among the native Scots pine sites in Scotland. To investigate 

whether differentiation in response to environmental variation has occurred in Scotland, a glasshouse-based commongarden trial of ~3,360 seedlings from 21 populations and 

84 open-pollinated families was established in 2007. At the 

beginning of the 2nd growing season, timing of bud flush 

showed evidence of genetic differentiation among populations, with those from cooler origins generally flushing earlier. Variation was also found among families within populations, suggesting that the trait is genetically controlled. 

Populations and families showed different levels of variability in this trait which could be partly due to variable levels 

of temporal climate fluctuation in different parts of Scotland. 

Chlorophyll fluorescence was used to examine drought response in three-year old seedlings from five populations on 

sites that experience contrasting levels of annual rainfall. It 

was found that the response was not related to rainfall, but 

possibly to more complex moisture variables that also take 

into account additional factors such as evaporation. Also, 

photosynthetic capacity in response to cold winter temperatures varied significantly among eight populations that were 

kept outdoors, and the largest reduction was seen in seedlings from the mildest, most maritime coastal site. The following spring, height growth and needle flush started earlier 

in seedlings from cooler locations. Earlier studies on genetic 

diversity of native pinewoods have shown high levels of selectively neutral variation in this predominantly outcrossing 

conifer, and a mating system analysis with a limited number 

of microsatellite markers supported this pattern. Together, 

these data suggest that despite significant historic population size decrease, environmental gradients have resulted in 

genetic differentiation among native pinewoods. In order to 

minimise the risk of planting poorly-adapted stock and to 

maximise the success of replanting programmes, it is important that the origins of planting stock are carefully considered in management guidelines for the species.

Type: Journal 

Geographic Area: Michoacán, Mexico

Keywords: Pinus oocarpa, provenances, altitudinal genetic 

variation, seed zoing, tree breeding, conservation, global warming, Mexico

Abstract: Pinus oocarpa has a large natural distribution in 

the sub-tropical forests of Mexico. Populations, however, 

are rapidly disappearing particularly in the Michoacán State 

as native forests are converted to avocado (Persea sp.) orchards. We investigated the patterning of genetic variation 

among P. oocarpa populations for quantitative traits along 

an altitudinal gradient by establishing a provenance/progeny 

test from wind-pollinated seeds collected along an altitudinal 

transect (1100–1500 m) near Uruapan, Michoacán, Mexico. 

Genetic variation was analyzed in relation to normalized climate records (temperature, precipitation, degree days > 58 

°C and annual moisture index) for the provenances and the 

test site for the contemporary climate and for climates projected for the decades beginning in 2030, 2060, and 2090. 

Estimates of future climates used output from the Canadian 

and Hadley General Circulation Models. Results of a field 

test suggested an altitudinal pattern of genetic differentiation 

in juvenile height among P. oocarpa populations. Seedlings 

from populations originating from lower altitudes tended to 

grow more than seedlings originating from populations at 

the higher altitudes. However, this trend abates at the lowest 

altitudinal limit of the species distribution, a probable conservative growth strategy for avoiding drought stress. Thus, 

the cline appeared to arise from selection along a climatic 

gradient reflecting in a moisture index (ratio of degree days 

to precipitation) and is dependent, therefore, on a balance 

between temperature and moisture. For guiding seed and 

seedling transfer in ecological restoration, conservation of 

genetic resources, tree breeding and mitigating the effects of 

global warming, we suggest guidelines based on delimitation of three altitudinal seed zones of about 200 m in breadth. 

Alternatively, one can limit transfer to three climatic zones 

of about 0.75 units of annual moisture index. Predictions of 

future climates indicate an average annual temperature increase of 3.8 °C by year 2090, and, judging from an increase 

of an annual moisture index of 26%, an increase in aridity 

by the end of the century. However, the more difficult period 

for adaptation of P. oocarpa populations to the new climate 

should be between 2030 and 2060, when the increase in 

aridity is expected to be most pronounced. Changes of this 

magnitude should alter the natural distribution of the species 

and would create an adaptation lag, as the adaptedness of 

extant populations deteriorates. Mitigating these effects will 

require seeds to be transferred upwards in altitude, perhaps 

as much as 150 m initially. 

Link: http://www.treesearch.fs.fed.us/pubs/27838

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planting maladapted seed stock, the patterns of environmental 

and adaptive genetic variation should be taken into account in 

the management of genetic resources in this species.

771. Salmela MJ, Cavers S, Wachowiak W, Cottrell 

JE, Iason GR, Ennos RA. 2010. Understanding the evolution of native pinewoods in Scotland will benefit their future management and conservation. Forestry 83:535-545.

Type: Journal 

Geographic Area: Scotland, United Kingdom

Compilers’ Keywords: Scots pine, Pinus sylvestris, evolutionary adaptation, seed zones, heavy exportation, timber production, phenotypic variability, common garden

Summary: Scots pine (Pinus sylvestris L.) is a foundation species in Scottish highland forests and a national icon. Due to 

heavy exploitation, the current native pinewood coverage represents a small fraction of the postglacial maximum. To reverse 

this decline, various schemes have been initiated to promote 

planting of new and expansion of old pinewoods. This includes 

the designation of seed zones for control of the remaining genetic resources. The zoning was based mainly on biochemical similarity among pinewoods but, by definition, neutral 

molecular markers do not reflect local phenotypic adaptation. 

Environmental variation within Scotland is substantial and it is 

not yet clear to what extent this has shaped patterns of adaptive differentiation among Scottish populations. Systematic, 

range-wide common environment trials can provide insights 

into the evolution of the native pinewoods, indicating how environment has influenced phenotypic variation and how variation is maintained. Careful design of such experiments can also 

provide data on the history and connectivity among populations, by molecular marker analysis. Together, phenotypic and 

molecular datasets from such trials can provide a robust basis 

for refining seed transfer guidelines for Scots pine in Scotland 

and should form the scientific basis for conservation action on 

this nationally important habitat.

772. Schmidtling RC, Myszewski JH, McDaniel CE. 

2005. Geographic variation in shortleaf pine (Pinus echinata

Mill.)—cortical monoterpenes. In: McKeand SE, Li B, editors. 

Proceedings of the 28th Southern Forest Tree Improvement 

Conference. Springfield (VA): The National Technical 

Information Service. p 15.

Type: Conference Paper

Geographic Area: Eastern USA

Keywords: shortleaf pine, Pinus echinata Mill., loblolly pine, 

Pinus taeda L., pond pine, Pinus serotina Michx., spruce pine, 

Pinus glabra Walt., terpenes, geographic variation, hybridization

Abstract: Cortical monoterpenes were assayed in bud tissue from 16 Southwide Southern Pine Seed Source Study 

(SSPSS) sources and from 6 seed orchard sources from across 

Link: https://www.era.lib.ed.ac.uk/bitstream/1842/5662/2/

Salmela2011.pdf

770. Salmela MJ, Cavers S, Cottrell JE, Iason GR, 

Ennos RA. 2011. Seasonal patterns of photochemical capacity and spring phenology reveal genetic differentiation among native Scots pine (Pinus sylvestris L.) populations in Scotland. Forest 

Ecology and Management 262:1020-1029.

Type: Journal 

Geographic Area: Scotland, United Kingdom

Keywords: adaptation, chlorophyll fluorescence, genetic differentiation, Scots pine, seasonal variation, spatial heterogeneity

Abstract: Environment-driven genetic differentiation among 

populations is a common feature among forest trees, and an 

understanding of how populations have adapted to their home 

site conditions is essential for management and conservation 

practices. In Scotland, 84 native Scots pine (Pinus sylvestris L.) 

woodlands are recognised by the Forestry Commission and 

they occupy highly diverse environments from the maritime 

west coast to continental sites in eastern Scotland. However, it 

is not known whether adaptations to local environments along 

sharp temperature and rainfall gradients have occurred in different populations and as a result, the seed transfer guidelines 

of the species are based only on data from isozymes and monoterpenes. In this study of an outdoor common garden trial, we 

used chlorophyll fluorescence to examine whether seedlings 

from 32 open-pollinated families and eight populations from 

sites experiencing contrasting annual temperature regimes 

differed in their response to variation in natural outdoor temperatures between September 2009 and May 2010. In addition, 

growth initiation in spring was recorded. Photochemical capacity at photosystem II Fv/Fm showed a distinct seasonal trend 

and remained at relatively high levels (~ 0.7) until November. 

Following a period of over 2 weeks with temperatures below 

or close to 0 °C, Fv/Fm started decreasing towards its minimum values recorded in early March when population means 

varied between 0.35 and 0.45. By early May and along with 

rising temperatures, photochemical capacity had recovered to 

the same level as observed in early November. Populations 

were found to respond differently to the cold period starting 

in December. The largest drop in photochemical capacity was 

observed in seedlings from a low-altitude population located 

in the maritime western Scotland, while in seedlings from 

higher altitude locations in the cooler eastern Scotland, the 

response was smaller. In March, the recovery of photochemical capacity was slowest in seedlings from the mildest and 

coolest sites. Evidence of adaptive genetic differentiation was 

also found in spring phenology. Initiation of shoot elongation 

and needle flush were earlier in families from higher altitudes 

(cooler areas), but population differences were not significant 

at the α = 0.05 level. These results suggest that adaptation to 

the spatially heterogeneous environment in Scotland has taken 

place in Scots pine and that in order to minimise the risk of 

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Geographic Area: Pacific Northwest USA

Compilers’ Keywords: Pinus ponderosa, Pseudotsuga menziesii, Abies grandis, Picea engelmannii, heat tolerance, survival 

analysis, forest management

Abstract: The heat tolerance of 2- to 4-week old seedlings of 

ponderosa pine (Pinus ponderosa Dougl. ex Laws), Douglasfir (Pseudotsuga menziesii (Mirb.) Franco), grand fir (Abies 

grandis (Dougl. ex D. Don) Lindl.), and Engelmann spruce 

(Picea engelmannii Parry ex Engelm.) was determined and 

compared by exposing them to various time-temperature 

combinations in a “dry water bath” apparatus. Isosurvival 

curves indicating the time-temperature combinations that resulted in approximately equal survival were constructed for 

each species. Small (2 to 4 °C) but significant differences in 

height tolerance were found among species with ponderosa 

pine having the greatest tolerance and Engelmann spruce the 

least. Douglas-fir and grand fir were generally intermediate 

in heat tolerance. 

775. Shaw NL. 2008. Native forbs and restoration technology. Native Plants Conference; Park City, UT. 74 p.

Type: Presentation

Geographic Area: Great Basin USA

Compilers’ Keywords: Great Basin Restoration Initiative, native plants, management, wildfire, plant materials, sagebrush

Compilers’ Summary: This is a presentation about the 

Great Basin Restoration Initiative. It is a collaborative program to promote the health and sustainability of the Great 

Basin for the public to use and enjoy. Components of the 

strategy include planning, inventory/assessment, implementation, monitoring/evaluation, and science. The presentation 

focuses on one part of the initiative - the Great Basin Native 

Plant Selection and Increase Project, which provides tools 

and support to native seed growers in meeting native seed 

demands. Research on native grass and forb species are 

highlighted, including cultural methods to establish them on 

sites. 

776. Shaw NL, Monsen SB. 1995. ‘Lassen’ antelope 

bitterbrush. In: Roundy BA, McArthur ED, Haley JS, Mann 

DK, compilers. Proceedings: Wildland Shrub and Arid Land 

Restoration Symposium. Ogden (UT): USDA Forest Service, 

Intermountain Research Station. General Technical Report INTGTR-315. p 364-371.

Type: Government Document

Geographic Area: North America

Compilers’ Keywords: antelope bitterbrush, Purshia tridentata, seed transfer guidelines

Abstract: ‘Lassen’ antelope bitterbrush is recommended 

for restoring depleted rangelands, burned areas, mined 

lands, and other disturbed sites in the Intermountain West. 

the natural range of the species, to examine geographic variation in shortleaf pine. Spruce pine and pond pine were also 

sampled. The results show geographic differences in all of the 

major terpenes. There was no north-south trend in any of the 

terpenes, but there was clinal variation in alpha pinene from 

west to east. One source, from New Jersey (SSPSSS) had very 

low alpha pinene and did not fit the trend, possibly because 

of hybridization with pitch pine. Some of the western sources 

had high limonene content, probably as a result of hybridization with loblolly pine, which has high limonene in western 

populations. Spruce pine had terpene levels similar to shortleaf 

pine, while pond pine had low alpha pinene and much higher 

limonene compared to shortleaf pine.

Link: http://www.treesearch.fs.fed.us/pubs/25247

773. Schuster WSF, Sandquist DR, Phillips SL, 

Ehleringer JR. 1994. High levels of genetic variation in 

populations of four dominant aridland plant species in Arizona. 

Journal of Arid Environments 27:159-167.

Type: Journal 

Geographic Area: Arizona, USA

Keywords: evolution, genetic diversity, genetic variation, isozymes, life expectancy, plant ecology, population genetics

Abstract: Intrapopulation genetic variation in four dominant, 

perennial plant species from the deserts of south-western North 

America was assessed and compared using isozyme techniques. 

In each of two localities, a warm desert and a cold desert environment, one population of the most common long-lived perennial was compared with a nearby population of a dominant, 

comparatively short-lived perennial. The warm desert species 

were Encelia farinosa and Larrea tridentata while Gutierrezia 

microcephala and Coleogyne ramosissima were examined at the 

cold desert site. All samples were electrophoresed and stained 

for 18 enzyme systems. Mean values for these four species 

were 0.173 for gene diversity, 99.6% for polymorphism, and 

2.02 for number of alleles per locus. These values are significantly greater than published means from recent survey of the 

plant isozyme literature, and are comparable to or higher than 

levels of genetic variation in other widespread plant species. 

Differences in level of variation among the species were not 

significant, thus failing to indicate any relationship between 

life expectancy and genetic variation. These results demonstrate that aridland plant populations can harbor high levels of 

genetic variation, and suggest that environmental heterogeneity may be important in the development and maintenance of 

this diversity.

Link: http://www.ehleringer.net/Jim/Publications/182.pdf

774. Seidel KW. 1986. Tolerance of seedlings of ponderosa 

pine, Douglas-fir, grand fir, and Engelmann spruce for high temperatures. Northwest Science 60:1-7.

Type: Journal 

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distance between sites was shown to be positively correlated 

with an increased difference in seed yield. Differences in 

size, growth habit, and leaf shape between paired habitats 

of origin within region and also between ‘‘ecotypes’’ were 

evident. However, there was no correlation between ecological distance and any of the measured traits. These findings 

suggest that: (1) Differences in phenotype and fecundity 

between geographically separated populations of L. corniculatus may be sufficient to lead to differences in survival and 

fitness when seeds are sown in a restoration environment. (2) 

Although it is important to consider geographical location, 

the choice of habitat within region is also important because 

phenotypic variation between ecotypes may have long-term 

consequences for plant persistence.

779. Sorensen FC, Mandel NL, Aagaard JE. 2001.

Role of selection versus historical isolation in racial differentiation of ponderosa pine in southern Oregon: an investigation of 

alternative hypotheses. Canadian Journal of Forest Research 

31:1127-1139.

Type: Journal 

Geographic Area: Oregon, USA

Compilers’ Keywords: Pinus ponderosa, cold hardiness, molecular marker, principal component analysis, climatic adaptation, physiography, common garden

Abstract: Continuous populations identified as Pacific and 

North Plateau races of ponderosa pine (Pinus ponderosa

P. Laws. ex C. Laws.) are parapatric along the crest of the 

Cascade Range in southern Oregon. A 3-year commongarden study of bud phenology and seedling vigor was 

performed to estimate the nature and magnitude of differentiation between races, to characterize the transition zone 

between them, and to relate responses between and within 

races to topography and climate. Principal component (PC) 

analyses identified two significant character complexes, PC-1 

(phenological traits) and PC-2 (size traits), that explained 

73% of the geographic race-related variation. The races were 

differentiated in two regards. First, PC-1 scores, which were 

highly correlated with frost-free season and summer–winter 

temperature differential, displayed a sharp discontinuity in 

the transition zone. Second, PC-2 scores were significantly 

correlated with physiographic and climatic variables in the 

North Plateau but not in the Pacific race, even though these 

variables had greater ranges in the latter. The data supported 

a narrow, adaptive transition between races for a complex 

of traits probably related to cold hardiness, and provided 

evidence that plant vigor traits were more closely adapted to 

environments in the North Plateau than in the Pacific region, 

possibly as a consequence of past climatic-stress selection 

in the former and competitive-stress selection in the latter.

Link: http://www.treesearch.fs.fed.us/pubs/4984 

Lassen was selected for its seedling vigor, upright growth 

habit, seed production, productivity, and retention of overwintering leaves. Selection trials indicate Lassen is adapted 

to antelope bitterbrush sites receiving 300 to 600 mm of precipitation at elevations ranging from 900 to 1,800 m. Best 

growth occurs on deep, coarse, well-drained, and neutral to 

slightly acidic soils. Productive Lassen plantings have been 

established on such sites in northeastern California, eastern 

Oregon, and central and southern Idaho. 

Link: http://www.treesearch.fs.fed.us/pubs/34717

777. Simpson DG, Binder WD, L’Hirondelle S. 

1999. Paper birch genecology and physiology. Journal of 

Sustainable Forestry 10:191-198.

Type: Journal 

Geographic Area: Western Canada

Compilers’ Keywords: Betula papyrifera, common garden, 

seed transfer guidelines, seed origin

Introduction: Forest managers in British Columbia increasingly manage some forests as mixed species stands that 

include paper birch (Betula papyrifera Marsh.), which regenerates naturally or can be planted. This genecology study, in 

association with ongoing tree improvement field studies, is 

aimed at developing seed transfer guidelines and identifying 

faster growing sources of paper birch for use in BC forests. 

Two of the objectives discussed here are to examine the effects of seed origin on (1) dormancy release (bud flushing), 

and (2) growth cessation and cold acclimation.

778. Smith BM, Diaz A, Daniels R, Winder L, 

Holland JM. 2009. Regional and ecotype traits in Lotus corniculatus L., with reference to restoration ecology. Restoration 

Ecology 17:12-23.

Type: Journal 

Geographic Area: United Kingdom

Keywords: ecological distance, geographical distance, local 

adaptation, seed origin

Abstract: To assess the relative importance of distance 

(geographical or ecological) as a consideration when collecting seeds for restoration projects, there is a need for more 

research on regional and ecotypic variation in a range of 

species. We used Lotus corniculatus L., a legume frequently 

included in grassland seed mixes, to investigate phenotypic 

variation between British accessions in a common garden 

experiment. The aim of this study was to investigate the 

effect of seed origin on plant phenotype and fecundity and 

to determine whether there was any evidence of correlation with distance (geographical or ecological) in populations grown from seed collected from different locations. 

Regional differences were detected in plant size, growth 

habit, pubescence, leaf shape, and fecundity. Geographical 

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forest of Western Australia. An analysis of molecular variance of 

66 polymorphic amplified fragment length polymorphism markers partitioned 92% of the total genetic variation within populations and 8% among populations, indicating relatively weak 

but statistically significant population genetic differentiation. 

Ordination of these genetic data showed marked west/east and 

north/south gradients. Pairwise population genetic dissimilarity was correlated with both geographic distance and environmental distance derived from five climate variables. However, 

partial Mantel test showed that the relationship between genetic 

and geographic distance was not independent of environmental 

distance, suggesting a non-neutral signature in these markers. 

Bayesian outlier analysis identified two markers, and spatial 

analysis methods tests identified highly significant associations 

between these two markers and three environmental variables. 

Frequency differences at these markers across populations suggested the possibility of climatically adapted provenances. The 

global significance value from analyses of similarities for these 

two markers correlated to a general provenance distance of 47 

km, in contrast to a threshold of 60 km for the complete dataset. Guidelines for seed sourcing that consider these population 

genetic data should lead to more effective ecological restoration 

with this species. 

782. Stonecypher RW, Piesch RF, Helland GG, 

Chapman JG, Reno HJ. 1996. Results from genetic 

tests of selected parents of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) in an applied tree improvement program.

Forest Science 42:1-37.

Type: Journal 

Geographic Area: Pacific Northwest USA

Keywords: genotype by environment interaction, genetic variation, stability, heritability, phenotypic selection

Abstract: Results from genetic tests, and genotype by environment interaction studies in six low elevation breeding 

zones of Weyerhaeuser Company’s Western Washington and 

Oregon Douglas-fir tree improvement program are summarized. Phenotypic selection in natural stands resulted in 

a 5% improvement in juvenile height over nonselect seed 

lots. Comparisons with nonselect sources, of offspring from 

parents in the top 50% performers in tests, indicated a 10% 

increase for the same trait. Seed produced in a 50% rogued 

seed orchard is thus expected to provide improved planting 

stock with a gain of 10% in juvenile height growth. Several 

select parents are producing offspring that are consistently 

performing in excess of 10% over nonselects. Estimates 

of breeding zone, breeding zone by location, and family 

by location interaction effects are small relative to family 

and planting location effects. Tests of families established 

on environmentally diverse sites indicate a striking lack of 

large family by planting location interaction. In tests showing statistically significant interactions, such interactions 

are caused by a relatively small number of families. We did 

780. St. Clair JB. 2006. Genetic variation in fall cold 

hardiness in coastal Douglas-fir in western Oregon and 

Washington. Canadian Journal of Botany 84:1110-1121.

Type: Journal 

Geographic Area: Pacific Northwest USA

Keywords: cold hardiness, genetic variation, adaptation, 

Pseudotsuga menziesii

Abstract: Genetic variation in fall cold damage in coastal 

Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco var. menziesii) 

was measured by exposing excised branches of seedlings from 

666 source locations grown in a common garden to freezing temperatures in a programmable freezer. Considerable variation was 

found among populations in fall cold hardiness of stems, needles, 

and buds compared with bud burst, bud set, and biomass growth 

after 2 years. Variation in fall cold hardiness was strongly correlated (r = 0.67) with cold-season temperatures of the source 

environment. Large population differences corresponding with 

environmental gradients are evidence that natural selection has 

been important in determining genetic variation in fall cold hardiness, much more so than in traits of bud burst (a surrogate for 

spring cold hardiness), bud set, and growth. Seed movement 

guidelines and breeding zones may be more restrictive when considering genetic variation in fall cold hardiness compared with 

growth, phenology, or spring cold hardiness. A regional stratification system based on ecoregions with latitudinal and elevational 

divisions, and roughly corresponding with breeding zones used 

in Oregon and Washington, appeared to be adequate for minimizing population differences within regions for growth and phenology, but perhaps not fall cold hardiness. Although cold hardiness 

varied among populations, within-population and within-region 

variation is sufficiently large that responses to natural or artificial 

selection may be readily achieved. 

Link: http://tafcc.forestry.oregonstate.edu/pdf/StClair_2006.pdf

781. Stingemore JA, Krauss SL. 2012. Genetic delineation of local provenance in Persoonia longifolia: implications 

for seed sourcing for ecological restoration. Restoration Ecology 

21:49-57.

Type: Journal 

Geographic Area: Australia

Keywords: AFLP, ANOSIM, landscape genomics, seed collection zone, spatial analysis method

Abstract: Restoration of diverse native plant communities typically requires the collection of large amounts of seed. Thus, practitioners often struggle to find adequate supplies near project sites 

and need to know from how far they can collect without compromising restoration success—how far does local provenance 

extend? We addressed this issue by assessing genetic variation 

within, and differentiation among, 12 potential seed source 

populations of Persoonia longifolia, a key component of the jarrah 
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784. Turesson G. 1922. The genotypical response of the 

plant species to the habitat. Hereditas 3:211-350.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: genomics, plant adaptation, genotype, 

phenotype, transplant experiments

Introduction: The study of the variability of the plant species 

in relation to its environment, or its habitat, might be pursued 

along two lines of research, viz. the study of the effect of various 

environmental factors upon the individual plant, and the study 

of the effect of these environmental factors upon the genotypical 

composition of a species-population in a certain habitat. The one 

line of research is primarily a study of the modifications of the 

plant, the other is manifestly a study of the hereditary variations. 

The two groups of variation have long ago been dealt with from 

the point of view of the environment, but while the experimental 

study of the former problem has received much attention the latter problem, the hereditary variation of wild plants in relation 

to habitat, has remained experimentally almost unattacked. In 

the absence of critical acquaintance with the different forms of 

a plant species met with in nature, much speculation is to the 

origin of adaptive structures is to be found in writings on ecology. The following discussion includes a number of notorious 

adaptive forms, and an attempt is made to ascertain whether the 

existence in nature of such forms is the result of an advantageous 

response on the part of the individual, or whether these forms 

are brought into existence through a genotypical response of the 

species-population to definite habitat conditions.

785. Vihera-Aarnio A. 2009. Effects of seed origin 

latitude on the timing of height growth cessation and field 

performance of silver birch [Dissertation]. Vantaa, Finland: 

University of Helsinki. 47 p.

Type: Dissertation

Geographic Area: Finland, Europe

Keywords: climatic adaptation, critical night length, moose 

browsing, provenance, seed transfer, stem quality, yield

Abstract: The aim of this thesis was to increase our knowledge 

about the effects of seed origin on the timing of height growth 

cessation and field performance of silver birch (Betula pendula 

Roth) from different latitudes, with special attention paid to the 

browsing damage by moose in young birch plantations. The effect of seed origin latitude and sowing time on timing of height 

growth cessation of first-year seedlings was studied in a greenhouse experiment with seven seed origins (lat. 58o - 67o N). 

Variation in critical night length (CNL) for 50% bud set within 

two latitudinally distant stands (60o and 67o N) was studied in 

three phytotron experiments. Browsing by moose on 5-11 yearold silver birch saplings from latitudinally different seed origins (53o - 67o N) was studied in a field experiment in southern 

Finland. Yield and stem quality of 22-year-old silver birch trees 

observe earlier budbreak and a higher spring frost susceptibility of Oregon sources established on Washington sites. 

We conclude that allocation and utilization of select families 

within Weyerhaeuser’s Oregon and Washington ownership 

should not be constrained by the currently defined breeding zone boundaries, but based on parental performance and 

stability for growth and adaptive traits in general. We further 

suggest that the necessity for maintaining separate breeding 

zones, within Washington and Oregon, in subsequent cycles 

of recurrent selection is questionable. Average individual 

tree heritability, from 65 6-parent disconnected diallels, is 

0.13 for age 6 and age 8 height. Dominance genetic variance 

is estimated to be one-half that of additive genetic variance 

for the same traits. The environmental designs and test establishment methods used resulted in acceptable levels of 

statistical precision for juvenile growth traits. Individual test 

least significant range values (LSR) resulted in significant 

family mean percent height differences of 8 to 26, and averaged 13% for diallels, and full-sib and half-sib tests.

783. Thomson AM, Crowe KA, Parker WH. 2010.

Optimal white spruce breeding zones for Ontario under current and future climates. Canadian Journal of Forest Research 

40:1576-1587.

Type: Journal 

Geographic Area: Ontario, Canada

Compilers’ Keywords: climate change, Picea glauca, range 

shifts, seed transfer guidelines

Abstract: Optimal breeding zones were developed for 

white spruce (Picea glauca (Moench) Voss) in Ontario under 

present and future climate conditions to examine potential 

shifts due to climate change. These zones were developed 

by (i) determining a set of candidate breeding zones based 

on the relationship between measured performance variables 

and climate and (ii) employing a decision support model to 

select subsets of breeding zones that maximize geographic 

coverage subject to a constraint on the maximum number 

of zones. Current optimal breeding zones were based on 

1961–1990 climate normals, and future breeding zones 

were based on three general circulation model (CGCM2, 

HADCM3, and CSIRO) predictions of 2041–2070 climate. 

Based on a maximum adaptive distance of 2.0 least significant difference values between sites within zones, 14 zones 

were required to cover the Ontario range of white spruce 

for the 1961–1990 data. Compared with breeding zones of 

other boreal-conifers, current optimal breeding zones for 

white spruce were quite large, spanning up to 38 latitude and 

108–128 longitude and indicating large distances of effective 

seed transfer. Of the three general circulation models used to 

simulate future climate, HADCM3 B2 and CGCM2 B2 predicted 2041–2070 breeding zones that largely coincide with 

1961–1990 zones. In contrast, CSIRO B2 indicated much 

narrower 2041–2070 breeding zones.
276 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

largely mediated by a positive association between stand basal 

area and tree species richness. Mean annual temperature and 

mean annual precipitation affected wood production and species 

richness directly. However, the direction and magnitude of the 

influence of climatic variables on wood production and species 

richness was not consistent, and vary dependent on forest type. 

Our analysis is the first to find a local scale positive relationship 

between tree species richness and tree wood production occurring across a continent. Our results strongly support incorporating the role of biodiversity in management and policy plans for 

forest carbon sequestration.

Link: http://www.ncbi.nlm.nih.gov/pubmed/23437038

787. Vitasse Y, Delzon S, Bresson CC, Michalet R, 

Kremer A. 2009. Altitudinal differentiation in growth and 

phenology among populations of temperate-zone tree species 

growing in a common garden. Canadian Journal of Forest 

Research 39:1259-1269.

Type: Journal 

Geographic Area: Southwestern France

Compilers’ Keywords: genetic variation, tree species, 

phenology, common garden, climate change, forest management 

planning

Abstract: The aim of the study was to determine whether there 

are genetic variations in growth and leaf phenology (flushing and 

senescence) among populations of six woody species (Abies alba 

Mill., Acer pseudoplatanus L., Fagus sylvatica L., Fraxinus excelsior L., Ilex aquifolium L., and Quercus petraea (Matt.) Liebl.) 

along altitudinal gradients, using a common-garden experiment. 

We found (i) significant differences in phenology and growth 

among provenances for most species and (ii) evidence that these 

among-population differences in phenology were related to the 

annual temperature at the provenance sites for ash, beech, and 

oak. It is noteworthy that along the same climatic gradient, species can exhibit opposing genetic clines: beech populations from 

high elevations flushed earlier than those from low elevations, 

whereas we observed the opposite trend for ash and oak. For 

most species, significant altitudinal clines for growth were also 

revealed. Finally, we highlighted the fact that both phenology 

timing and growth rate were highly consistent from year to year. 

The results demonstrated that despite the proximity of the populations in their natural area, differences in altitude led to genetic 

differentiation in their phenology and growth. These adaptive 

capacities acting along a natural climatic gradient could allow 

populations to cope with current climate change.

Link: http://www.nrcresearchpress.com/doi/abs/10.1139/

X09-054#.VTndbpgcRLM

788. Viveros-Viveros H, Sáenz-Romero C, VargasHernández JJ, López-Upton J, Ramírez-Valverde 

G, Santacruz-Varela A. 2009. Altitudinal genetic 

variation in Pinus hartwegii Lindl. I: Height growth, shoot 

of Baltic, Finnish and Russian origin (54o - 63o N) and the effect 

of latitudinal seed transfers were studied in two provenance trials 

at Tuusula, southern and Viitasaari, central Finland. The timing 

of height growth cessation depended systematically on latitude of 

seed origin and sowing date. The more northern the seed origin, 

the earlier the growth cessation and the shorter the growth period. 

Later sowing dates delayed growth cessation but also shortened 

the growth period. The mean CNL of the southern ecotype was 

longer, 6.3 ± 0.2 h (95 % confidence interval), than that of the 

northern ecotype, 3.1 ± 0.3 h. Within-ecotype variance of the CNL 

was higher in the northern ecotype (0.484 h2) than in the southern 

ecotype (0.150 h2). Browsing by moose decreased with increasing latitude of seed origin and sapling height. Origins transferred 

from more southern latitudes were more heavily browsed than 

the more northern native ones. Southern Finnish seed origins 

produced the highest volume per unit area in central Finland (lat. 

63o11’ N). Estonian and north Latvian stand seed origins, and the 

southern Finnish plus tree origins, were the most productive ones 

in southern Finland (lat. 60o21’ N). The relationship of both survival and stem volume/ha to the latitudinal seed transfer distance 

was curvilinear. Volume was increased by transferring seed from 

ca. 2 degrees of latitude from the south. A longer transfer from the 

south, and transfer from the north, decreased the yield. The proportion of trees with a stem defect increased linearly in relation to 

the latitudinal seed transfer distance from the south.

Link: http://www.metla.eu/dissertationes/df87.pdf

786. Vila M, Carrillo-Gavilan A, Vayreda J, Bugmann 

H, Fridman J, Grodzki W, Haase J, Kunstler G, 

Schelhaas M, Trasobares A. 2013. Disentangling 

biodiversity and climatic determinants of wood production. PLoS 

ONE 8:e53530.

Type: Journal 

Geographic Area: Europe

Compilers’ Keywords: biodiversity, climate change, wood 

production, tree growth, carbon sequestration, species richness

Abstract: Despite empirical support for an increase in ecosystem productivity with species diversity in synthetic systems, there 

is ample evidence that this relationship is dependent on environmental characteristics, especially in structurally more complex 

natural systems. Empirical support for this relationship in forests 

is urgently needed, as these ecosystems play an important role in 

carbon sequestration. We tested whether tree wood production 

is positively related to tree species richness while controlling 

for climatic factors, by analyzing 55265 forest inventory plots 

in 11 forest types across five European countries. On average, 

wood production was 24% higher in mixed than in monospecific forests. Taken alone, wood production was enhanced with 

increasing tree species richness in almost all forest types. In some 

forests, wood production was also greater with increasing numbers of tree types. Structural Equation Modeling indicated that 

the increase in wood production with tree species richness was 

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Endocronartium harknessii (J.P. Moore) Y. Hiratsuka. Greater 

foliar concentrations of lignin, tannins, and some phenolics were 

associated with increased resistance to single or multiple foliar 

pathogens. Bark secondary metabolites levels were generally 

unassociated with resistance to bark diseases. Provenances appearing to originate in ecosystems where lodgepole pine are not 

the dominant species generally were more susceptible to foliar 

diseases and had less foliar defense associated compounds than 

trees from areas where pines were the dominant species, yet clear 

trends proved to be elusive. Regardless, pine provenances with 

greater foliar levels of identified defense-associated compounds 

should be preferred seed sources for replanting forests in areas 

in which foliar disease is expected to be increasingly prevalent.

790. Wang JR, Hawkins CDB, Letchford T. 1998.

Photosynthesis, water and nitrogen use efficiencies of four paper 

birch (Betula papyrifera) populations grown under different soil 

moisture and nutrient regimes. Forest Ecology and Management 

112:233-244.

Type: Journal 

Geographic Area: Western Canada

Keywords: Betula papyrifera, paper birch, nitrogen use efficiency, photosynthesis, population, water use efficiency, 

silviculture

Abstract: A greenhouse experiment was conducted to determine how four populations of paper birch (Betula papyrifera Marsh.), adapted to particular temperature, water and 

photoperiod regimes responded physiologically to different 

soil water and nutrient regimes. Seedlings of each of the four 

populations (Eaglet, LeeCreek, Porcupine and Skeena) from 

British Columbia were planted in pots and subjected to high 

and low water and nitrogen regimes for 4 months. Net photosynthesis (A), stomatal conductance (gs), water use efficiency (WUE) and nitrogen use efficiency (NUE) were measured and compared among treatments. Trade-offs between 

WUE and NUE were examined. Soil moisture was the major 

factor affecting A for all the populations. All populations 

showed relatively high A in the high water and high-nitrogen 

(HWHN) treatment, ranging from 8.5 to 9.9 mmol CO2 m -2

s -1. Photosynthesis of trees in the low-water and high-nitrogen (LWHN) treatment was about 45% of that in the HWHN 

treatment. There was a linear increase in A as gs increased in 

all populations. A decreased with increasing vapor pressure 

deficit (VPD). The populations demonstrated significantly 

different relationships between A and gs, and VPD, with the 

LeeCreek population being the most conservative. This may 

provide a physiological basis for provenance selection and 

seed transfer. Eaglet and LeeCreek populations had lower 

WUE at the same level of NUE compared with Porcupine 

and Skeena populations. Porcupine and Skeena populations 

may be more suitable for moist sites due to their greater capacity to sequester water quickly, while the LeeCreek and 

phenology, and frost damage in seedlings. Forest Ecology 

and Management 257:836-842.

Type: Journal 

Geographic Area: Michoacán, Mexico

Keywords: altitudinal variation, genetic variation, population 

differentiation, frost damage, grass-stage, assisted migration, 

global warming, seed zones, Pinus hartwegii, Michoacán

Abstract: The altitudinal pattern of genetic variation in Pinus 

hartwegii Lindl. populations was explored for seedling height 

growth, frost damage, grass-stage, and phenological stage of the 

terminal shoot. A provenance test was conducted with open-pollinated seed from 13 populations collected along an altitudinal 

transect (3000–3600 m) at the National Park Pico de Tancítaro, 

Michoacán, center-west Mexico. Height growth of seedlings in 

a nursery was assessed at seven and 18 months of age. Frost 

damage at -15 °C was evaluated in laboratory at 18 months 

of age; proportion of plants that had left grass-stage and stage 

of shoot development was assessed at the age of 22 months. 

Significant differences among provenances (P < 0.0001) were 

detected for all of the evaluated characters. The variation among 

populations was structured as a moderate altitudinal cline, with 

populations from lower altitudes showing larger height growth 

in seedlings, larger proportion of frost damages, fewer seedlings in grass-stage and more seedlings with developed shoot, 

whereas in populations from higher altitudes, seedlings exhibiting shorter plant height, lower percentages of frost damage, 

more seedlings with unbroken grass-stage, and fewer seedlings 

with advanced shoot development were displayed. Options for 

seed and seedling movement along the altitudinal gradient are 

discussed under the scope of reforestation, aiming at ecological 

restoration, conservation of forest genetic resources, and assisted 

migration considering global warming. We suggest delineation 

of two latitudinal seed zones (Zone I: 3000–3350 m; Zone II: 

3350–3700 m).

789. Wallis CM, Reich RW, Lewis KJ, Huber DPW. 

2010. Lodgepole pine provenances differ in chemical defense 

capacities against foliage and stem diseases. Canadian Journal of 

Forest Research 40:2333-2344.

Type: Journal 

Geographic Area: Western Canada

Compilers’ Keywords: Pinus contorta, provenance test

Abstract: Maximization of lodgepole pine (Pinus contorta

Douglas ex Louden var. latifolia Engelm. ex S. Watson) growth 

in a future climate with increased pest activity requires an understanding of the natural variability of quantitative resistance 

to disease. Foliar and bark secondary metabolites from different lodgepole pine provenances (populations) were quantified and correlated with severity of foliar diseases caused by 

Lophodermella spp. (Lophodermella concolor (Dearn.) Darker 

or Lophodermella montivaga Petre.) or Elytroderma deformans

(Wier) Darker and bark diseases caused by Elytroderma or 

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278 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

be the best site-adapted material and the use of non-local 

material, or less `fit’ ecotypes may result in establishment 

failure, long-term mortality or disruption to the surrounding 

ecosystem. No guidelines for the definition of a provenance 

currently exist for any Australian native grass, providing 

a major obstacle for the broad-scale use of Australian native grasses. Understanding the evolutionary forces such as 

natural selection, gene flow and genetic drift as well as the 

factors that operate to drive these forces, can provide clues 

to the definition of a provenance boundary. If native grasses 

are to retain their adaptive and low input advantages over 

the use of exotic species then understanding the issues of 

provenance provides the key to their successful and widespread use.

Link: http://www.regional.org.au/au/stipa/papers/stipa2001-09.htm

793. Weißhuhn K, Auge H, Prati D. 2011.

Geographic variation in the response to drought in nine grassland species. Basic and Applied Ecology 12:21-28.

Type: Journal 

Geographic Area: Europe

Keywords: intraspecific variability, climate change, root allocation, reproductive allocation, specific leaf area

Abstract: Summer precipitation is expected to decrease in 

Europe due to climate change, which in turn will increase 

the risk for plants to suffer from drought. To maintain ecosystem functions in grasslands under the conditions of climate change, drought adapted varieties of plant species may 

be needed. With our experiment, we bridge the gap between 

single species experiments and field experiments with whole 

plant communities under proposed scenarios of climate 

change. In a greenhouse experiment, we investigated the 

drought response of nine common grassland species which 

are normally utilized for restoration of mesic grasslands. We 

chose five European provenances of each species to test for 

a consistent, provenance specific drought response. Plants of 

all nine species were negatively affected by drought with respect to their biomass production, and consistently allocated 

less resources to roots than control plants. On the other hand, 

reproductive allocation increased under drought conditions 

in some species whereas no response to drought could be detected in specific leaf area. In contrast to our expectation, we 

found little evidence for intraspecific variation in response 

to simulated drought. However, we found a clear correlation between responsiveness in root allocation, and climatic 

conditions at the provenance’s origin. This pattern became 

evident only when all species were analysed together, highlighting the advantage of multi species experiments.

Link: http://www.botany.unibe.ch/planteco/abstr_repr/

BAAE_12_21.pdf

Eaglet populations may be more suitable for drier interior 

sites where long drought periods can occur.

791. Ward K, Gisler M, Fiegener R, Young A. 2008.

The Willamette Valley Seed Increase Program: developing 

genetically diverse germplasm using an ecoregion approach. 

Native Plants Journal 9:334-350.

Type: Journal 

Geographic Area: Oregon, USA

Keywords: native plant material development, prairies, 

restoration genetics, seed transfer zones, domestication selection, Diversity Enhancement Block design

Abstract: The goal of the Institute for Applied Ecology’s 

Willamette Valley Seed Increase Program is to develop a supply of ecologically appropriate, genetically diverse native plant 

material for restoration of prairie ecosystems in the Willamette 

Valley. In creating restoration germplasm we seek to maximize 

genetic diversity while simultaneously protecting genetic integrity of extant native populations. In the absence of genetic data to 

guide appropriate movement of native seeds, we are testing the 

use of an ecoregion approach using a variety of research techniques. We collected seeds, defined preliminary seed transfer 

zones, and planted seed increase fields for each of 21 historically 

widespread, common species. We captured spatial and temporal 

genetic diversity by sampling from many populations per species over a 3-y period. Seed zone boundaries for each species 

were drawn at the scale of the ecoregion or smaller, depending 

on life history characteristics and potential for adverse genetic 

effects of translocation. To minimize loss of diversity through 

domestication selection, we planted increase fields using a novel 

design, the Diversity Enhancement Block. Seedlots from populations with different phenology or from different areas within 

the ecoregion were planted in separate adjacent blocks. This 

design allows harvest of each block separately as seeds mature, while still permitting plants from different regions of the 

valley to cross pollinate and to produce crop seeds with maximum genetic diversity. All of our production fields have been 

entered into the Oregon Seed Certification Service Pre-Variety 

Germplasm program. We are looking for partners to participate 

in a buyer’s cooperative.

792. Waters CM. 2001. Guiding the delimitation of local 

provenance for Australian native grasses. In: Second National 

Conference of the Native Grasses Association. New South 

Wales, Australia: Stipa Native Grasses Association, Inc. 1 p 

Type: Conference Paper

Geographic Area: Australia

Keywords: seed source, transfer guidelines, restoration, local 

adaptation

Abstract: The provenance of a seed source not only describes its point of collection but can also reflect its evolutionary development. Here, local provenance material may 

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scale. Multi-locus analyses reveal the converse: canonical correlation analysis of individual, uniformly-spaced genotypes 

describe statistically-significant, complex patterns with geography. Multi-locus scores in four major species. Abies concolor, 

Pinus lambertiana, P. ponderosa, and Pseudotsuga menziesii, of 

the mixed conifer forest in the Sierra Nevada correlate 0.40 or 

greater with the first canonical vector of a geographical trend 

surface equation. The different species follow similar patterns 

by latitude and elevation. In contrast with patterns in the Sierra 

Nevada, large-scale differentiation is weak (R2 < 0.20) among 

populations of Pseudotsuga menziesii in the Coast Ranges 

and Siskiyou Mountains of northern California and southern 

Oregon, where differentiation may be local. For the purpose 

of forming zones, we subdivided scores of the first two to four 

canonical vectors into groups and plotted them as multidimensional contour intervals. Reclassification by discriminant 

analysis serves as an approximate guide to transfer risks within 

and among these groups.

Link: http://www.treesearch.fs.fed.us/pubs/32853

796. Wilson BL, Darris DC, Fiegener R, Johnson R, 

Horning ME, Kuykendall K. 2008. Seed transfer zones 

for a native grass Festuca roemeri: genecological evidence. Native 

Plants Journal 9:287-303.

Type: Journal 

Geographic Area: Western USA

Keywords: common-garden study, habitat restoration, inbreeding depression, conservation genetics

Abstract: A common-garden study of Festuca roemeri (Pavlick) 

E. B. Alexeev (Poaceae) revealed substantial genetic variation 

within and among 47 populations from throughout its range in 

the Pacific Northwest, U.S., for growth, fitness, phenological, 

and morphological traits. Using climatic and physiographic 

variables, genetic patterns over the landscape were examined 

through principal component and regression analysis. Elevation 

and latitude of the seed source, and to a lesser extent temperature and precipitation, explained a significant proportion of the 

genetic variation, suggesting that observed variation was associated with adaptation to local environments. Most plants from the 

Willamette Valley exhibited poor growth and survival, perhaps 

due to inbreeding. Festuca roemeri variation clustered into seed 

transfer zones corresponding to Level III ecoregions, and one 

zone was further subdivided. High-elevation populations separated from lower-elevation populations but did not cluster into 

a single seed zone. Seed transfer zones reported here provide a 

guide for plant community restoration efforts using this species.

Link: http://www.treesearch.fs.fed.us/pubs/35452

797. Wu HX, Ying CC, Ju H-B. 2005. Predicting site productivity and pest hazard in lodgepole pine using biogeoclimatic 

system and geographic variables in British Columbia. Annals of 

Forest Science 62:31-42.

794. Weng YH, Parker WH. 2008. Adaptive variation 

in fall cold hardiness of aspen from northwestern Ontario. Tree 

Physiology 28:143-150.

Type: Journal 

Geographic Area: Ontario, Canada

Keywords: chlorophyll fluorescence, electrical conductivity, 

Populus tremuloides, principal component analysis, provenance 

trial, visual scoring

Abstract: We investigated adaptive variation in fall cold hardiness development based on the electrical conductivity of tissue 

diffusates (EC) among 20 aspen provenances from northwestern 

Ontario. Provenance accounted for over 40% of the total variation 

in cold injury for seven dates from September through November 

in three provenance trials. Principal component analysis was performed to summarize the combinations of results for all sampling 

sites, dates and temperatures (traits). Principal component (PC)-1 

represented fully developed cold hardiness differences among 

provenances; PC-2 represented differences in the timing of the 

onset of cold hardiness development; and PC-3 represented a 

site-related difference in cold hardiness development. Heat sum 

in early summer and late summer precipitation together were 

the best predictors of absolute degree of cold hardiness (PC-1), 

whereas temperatures for mid- to late summer were best for predicting onset of cold hardiness development (PC-2). In a second 

study, we assessed the efficacy of chlorophyll fluorescence (CF) 

as a simpler technique for determining the cold hardiness of aspen stem samples. Fall cold hardiness of stem samples of 12 of 

the original 20 provenances was estimated by CF, and the results 

were evaluated by a visual scoring (VS) method. Correlations between EC and CF measurements from the two studies were moderately strong based on the extent of cold hardiness in October 

of each year, but were negative for September dates because of a 

later onset of cold hardiness in the EC study year. Although the 

EC and CF methods gave similar cold hardiness values for stem 

samples from 12 provenances, the CF method may be preferred 

to the EC or VS method for species with chlorophyllous stems 

because of its greater ease of use.

Link: http://treephys.oxfordjournals.org/content/28/1/143.

full.pdf

795. Westfall RD, Conkle MT. 1992. Allozyme markers 

in breeding zone designation. New Forests 6:279-309.

Type: Journal 

Geographic Area: Western USA

Keywords: geographic variation, multivariate analysis, trendsurface analysis, transfer risk

Abstract: Early studies of allozyme variation in plant populations suggested that allelic frequencies in some loci vary by 

geography. Since then, the expectation that allozymes might 

be useful in describing geographic patterns has generally not 

been borne out by single locus analyses, except on the broadest 

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grew tallest. The remaining characters were only investigated 

10 years after outplanting. Among-provenance variation in mortality, frost damage, and stem defects was nonsignificant when 

two provenances from the southern end of the natural range 

were removed from the analysis. Resistance to needle disease 

(Uredinopsis longimucronata Faull) was significantly different 

among provenances and decreased with elevation. Geographic 

variation in the proportion of trees with lammas growth was also 

significant but did not display any apparent pattern. Variation 

patterns were similar at the four testing sites but among-site 

differences in the average performance were highly significant 

for all the traits investigated. Eastern Vancouver Island, British 

Columbia, and northeastern Olympia Peninsula, Washington, 

are recommended as primary areas of seed source for reforestation in the Vancouver forest region. 

799. Yang J, Pedlar JH, McKenney DW, Weersink 

A. 2015. The development of universal response functions to 

facilitate climate-smart regeneration of black spruce and white 

pine in Ontario, Canada. Forest Ecology and Management 

339:34-43.

Type: Journal 

Geographic Area: Ontario, Canada

Keywords: climate change, seed deployment, seed procurement, universal response function, black spruce, white pine 

blister rust

Abstract: Climate change is expected to impact forest growth 

and survival as tree populations experience climates to which 

they are not well adapted. However, forest regeneration efforts represent opportunities to introduce genetic material 

on the landscape that is well matched to future climates. We 

estimated universal response functions (URF) for two economically and ecologically important tree species in eastern 

Canada – black spruce (Picea mariana) and eastern white pine 

(Pinus strobus). The URF approach models tree growth as a 

function of both the planting site climate (i.e., environmental 

effects) and seed source climate (i.e., genetic effects), allowing the growth of any seed source to be estimated at any location. Both species exhibited a relatively weak genetic effect, 

suggesting modest potential for seed movements to mitigate 

climate change impacts. The optimal provenance for any given 

location was not local, but rather located in environments that 

were approximately 1.5 °C warmer than the planting site; this 

shift is consistent in both direction and magnitude with that 

expected due to climate warming since the start of the industrial revolution. Both species exhibited a strong environmental 

effect with clear evidence for a central climatic optimum. As 

a result, climate warming is expected to enhance black spruce 

growth at sites with mean annual temperature colder than approximately 4.5 °C and white pine growth at sites colder than 

11 °C. Thus, Ontario white pine populations may benefit from 

the climate changes projected to occur in the province over the 

next 50–100 years. For black spruce, populations in southern 

Type: Journal 

Geographic Area: Western Canada

Keywords: site productivity, pest hazard, western gall rust, 

needle cast, terminal weevil, ecological system

Abstract: A series of 60 lodgepole pine provenance tests was 

planted throughout the interior of British Columbia in 1974 to 

predict productivity and pest hazard based on ecological classification and geographical variables. These 60 tests cover eight 

biogeoclimatic zones and 25 subzones within the biogeoclimatic 

ecosystem classification (BEC) in British Columbia. Ten provenances are common among 60 provenances tested at each site. 

Mean height (20-year) was measured at 57 of the 60 sites, incidence of western gall rust assessed at 56 sites, terminal weevil 

at 49 sites, and needle cast at 50 sites. There is large site-to-site 

variation in all traits. Geographic models using latitude, longitude 

and elevation of test site location as predictors explained 47%, 

35%, 33%, 27% and 8% of site variation for height, survival, 

incidence of needle cast, terminal weevil and western gall rust, 

respectively. BEC zones accounted for about the same amount 

of the site variation as geographic models, suggesting both accounted for the effect of site environments relating mainly to 

temperature and precipitation. Within BEC zones, site variation 

in height seems to be related to subzones associated with moisture gradient, but not temperature. Sites in the moist, mild ICH 

subzone and the dry, cool MS subzone along the southern Rocky 

Mt. Trench represent the best forest land for intensive silviculture of lodgepole pine, being highly productive with low pest 

hazard except needle cast. These sites are followed in productivity by sites across the vast interior stretching from the Skeena/

Bulkley river basin in the northwest (moist SBS subzone), to the 

interior wetbelt on Shuswap Quesnel Highland (moist, cool ICH 

subzone), and the Thompson Plateau in the southern interior, 

where lodgepole pine grew well with relatively low pest hazard 

at most sites.

798. Xie CY, Ying CC. 1993. Geographic variation of 

grand fir (Abies grandis) in the Pacific coast region: 10-year results 

from a provenance trial. Canadian Journal of Forest Research 

23:1065-1072.

Type: Journal 

Geographic Area: Western North America

Compilers’ Keywords: reforestation, geographic differentiation, coastal forests, seed source

Abstract: The performance of 23 grand fir (Abies grandis 

(Dougl.) Lindl.) seed sources representing the coastal range 

of the species was examined with respect to height, mortality, 

frost damage, stem defects, needle disease susceptibility, and 

lammas growth at four sites in the Vancouver forest region of 

British Columbia. Variation in height was highly significant 

among provenances and showed discernible patterns at all 

ages surveyed (1, 3, 6, 7, 8, 9, and 10 years from outplanting). 

Provenances from northern, coastal, and low-elevation regions 

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Abstract: Thirty provenances of lodgepole pine (Pinus contorta Dougl.) test shock raised at two nurseries, Cowichan 

Lake (coastal British Columbia) and RE Rock (interior British 

Columbia), were assessed in two 15-year field trials. Analyses 

indicated three broad geographic regions of genetic differentiation in British Columbia: coast, coast–interior transition, and 

central and southern interior. Provenance elevation was found to 

have a strong influence on growth. The results suggest that the 

present seed transfer guidelines for lodgepole pine in the interior 

region of this province are conservative enough to prevent the 

use of maladapted seed sources. Nursery effect declined over a 

period of 15 years, while provenance differences increased with 

the age of the trials. Interactions between provenances and sites 

also increased after age 9. This suggests that the effect due to 

seedling culture and environment in the nursery is short-term 

relative to the influence of the genetic component. Nursery 

growth was generally not a good predictor of provenance field 

performance. 

802. Zhang JW, Feng Z, Cregg BM, Schumann CM. 

1997. Carbon isotopic composition, gas exchange, and growth 

of three populations of ponderosa pine differing in drought 

tolerance. Tree Physiology 17:461-466.

Type: Journal 

Geographic Area: Montana, South Dakota, USA

Keywords: carbon isotope ratio, net photosynthesis, 

Pinus ponderosa, relative growth rate, stomatal conductance, 

water-use efficiency, xylem pressure potential

Abstract: Effects of water supply on gas exchange, carbon 

isotopic composition, and relative growth rate were compared 

among seedlings from three populations of ponderosa pine 

(Pinus ponderosa Dougl. ex Laws.) grown in a controlled environment chamber. The three populations were chosen to represent high, moderate and low drought tolerance. There was no 

indication that drought tolerance was related to high water use 

efficiency. Populations differed (P < 0.05) in relative growth 

rate (RGR), but did not differ (P > 0.10) in gas exchange variables or carbon isotope ratio (δ13C). Well-watered seedlings had 

significantly higher RGR, xylem pressure potential (Yxpp), net 

photosynthesis (A), stomatal conductance to water vapor (g), 

and lower δ13C and instantaneous water-use efficiency than 

water-stressed seedlings. With decreasing Yxpp, A decreased 

linearly, whereas g decreased exponentially. Seedlings of the 

highly drought-tolerant population were more sensitive to water 

availability than seedlings from the other populations; they used 

water quickly when water was available, but closed their stomata in response to water stress. We conclude that, in ponderosa 

pine, the drought avoidance mechanism is more important for 

survival and growth in arid and semiarid environments than the 

efficient use of water.

Link: http://www4.nau.edu/direnet/publications/

publications_z/files/J_W._Zhang.pdf

and central Ontario may experience suboptimal climate conditions over the mid-to-end of the current century. Despite the 

relatively weak genetic effect, climate-smart seed movements 

could play a role in maintaining the productivity and genetic 

diversity of black spruce in these areas. While the URF approach has limitations, this work demonstrates its potential for 

informing regeneration decisions under climate change.

Link: http://www.sciencedirect.com/science/article/pii/

S0378112714007087

800. Ying CC. 1997. Effects of site, provenance, and 

provenance and site interaction in Sitka spruce in coastal British 

Columbia. Forest Genetics 4:99-112.

Type: Journal 

Geographic Area: Pacific Northwest, USA, Canada

Keywords: Picea sitchensis, site environment, provenance, interaction, growth

Abstract: The IUFRO International Sitka spruce (Picea sitchensis) provenance experiments were planted at eight sites in 

Coastal British Columbia in 1975. The tests were planted in two 

series, with four sites in each series and 10 provenances at each 

test. A total of 14 provenances were tested with 6 provenances 

common to both series. Total height and survival were recorded 

at 3, 6, 10, 15 and 20 years after planting, and diameter at ages 

10, 15 and 20 at most tests. Test sites accounted for about twothirds of the total variation. Size of provenance variance components was different between the two site series. Interaction between provenance and site was high and statistically significant 

in both series. Site climate and attack by the white pine weevil 

(Pissodes strobi) were the major determinants of site productivity and also the main causes of the interactions. Latitudinal pattern in provenance variation was eminent at most sites at early 

ages and maintained throughout at sites with strong maritime 

climate and free of weevil attack; at harsh northern and inland 

sites, geographic pattern of provenance variation shifted toward 

longitudinal (coast - inland) at later ages, and the harsher the site 

environment the earlier the shift. Practical implication in terms 

of seed transfer of the fast growing southern coastal sources and 

weevil resistant provenances in reforestation in coastal British 

Columbia was discussed.

Link: http://www.tuzvo.sk/files/fg/volumes/1997/FG04-

2_099-112.pdf

801. Ying CC, Thompson C, Herring L. 1989.

Geographic variation, nursery effects, and early selection in lodgepole pine. Canadian Journal of Forest Research 19:832-841.

Type: Journal 

Geographic Area: Western Canada

Compilers’ Keywords: genetic variation, provenance, 

common garden, Pinus contorta, seed transfer guidelines, 

maladaptation

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803. Adair R, Johnson RC, Hellier B, Kaiser WJ. 

2006. Collecting tapertip onion (Allium acuminatum Hook.) in 

the Great Basin using traditional and GIS methods. Native Plants 

Journal 7:141-148.

Type: Journal 

Geographic Area: Great Basin USA

Keywords: National Plant Germplasm System, georeferencing, field sampling, in situ conservation

Abstract: As part of a biodiversity assessment and germplasm 

conservation project, we collected 55 populations of tapertip 

onion (Allium acuminatum Hook. [Alliaceae]) throughout 

the Great Basin region of the U.S. (Idaho, Oregon, Nevada). 

Population data from field observations and herbarium specimens were entered into a geographic information system 

(GIS) to facilitate efficient sampling strategies and maximize 

the number and diversity of ecoregions represented. Although 

limited by the quality of source data, GIS-based population location information greatly aided in the organization and time 

management of the field collection. We collected, measured, 

and described bulbs. The bulbs will be used in common garden studies and genetic research to evaluate species diversity 

throughout the region, and develop in situ germplasm conservation sites for future native rangeland restoration efforts. 

Ex situ collections will also be entered into the National Plant 

Germplasm System and be available for research, ecological 

restoration efforts, and germplasm enhancement.

Link: http://www.nativeplantnetwork.org/Content/

Articles/7-2NPJ141-148.pdf

804. Breed MF, Stead MG, Ottewell KM, Gardner 

MG, Lowe AJ. 2013. Which provenance and where? Seed 

sourcing strategies for revegetation in a changing environment.

Conservation Genetics 14:1-10.

Type: Journal 

Geographic Area: Global

Keywords: climate change, habitat fragmentation, inbreeding, 

local adaptation, outbreeding depression, plant genetic resources, revegetation

Abstract: Revegetation is one practical application of science 

that should ideally aim to combine ecology with evolution to 

maximise biodiversity and ecosystem outcomes. The strict 

use of locally sourced seed in revegetation programs is widespread and is based on the expectation that populations are locally adapted. This practice does not fully integrate two global 

drivers of ecosystem change and biodiversity loss: habitat 

General (583–616)—reviews, history, 

summaries, justification

Research (617–802)—genetic studies, tree 

improvement, climate change

Strategies (803–827)—framework, delineation, modelling

Resources (828–877)—guidelines, zones, 

maps, tools, software

Transfer Guidelines

and Zones

CHAPTER 4 • TRANSFER GUIDELINES AND ZONES • STRATEGIES

#. Author(s), Date. Title. Source.

Type: 

Geographic Area: 

Keywords: 

Description:

Link:

location of study

keywords written by the 

author(s) or provided by the 

compilers

ID number and

information

abstract, summary, or 

portions of the introduction 

or conclusion written by the 

author(s) or provided by the 

compilers

book, book section, conference 

paper, dissertation, government 

document, journal article, 

newsletter, presentation, report, 

thesis or web page

internet address of reference

Each reference contains the following:
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 283

806. Crowe KA, Parker WH. 2005. Provisional breeding zone determination modeled as a maximal covering location 

problem. Canadian Journal of Forest Research 35:1173-1182.

Type: Journal 

Geographic Area: Central Canada

Compilers’ Keywords: seed zone, jack pine, modeling, focal 

point seed zone, common garden, Pinus banksiana, delineate 

breeding zones

Abstract: This study was a first attempt to model the problem of delineating breeding zones as a maximal covering 

location problem. The method involves two steps. First, a 

comprehensive set of candidate breeding zones is generated 

for a region using the focal point seed zone method. This 

method allows for control over the adaptive difference of 

genetic material within each zone. A grid of points is used 

to create the set of candidate breeding zones: one zone per 

point. Next, candidate zones are entered into a maximal covering location model formulated to suit this problem. The 

objective of this model is to select a subset of candidate 

zones that maximally covers the area of the region, given a 

limit on the number of zones to be selected and on the adaptive dissimilarity allowed within zones. Through use of this 

method, decision-makers can gain insight into how many 

breeding zones are needed to cover the region. Using different inputs from the focal point seed zone method, it is also 

possible to explore the trade-offs between the quantity and 

the quality of breeding zones. The method was tested on data 

from a series of jack pine (Pinus banksiana Lamb.) common 

garden trials of 102 seed sources from northwestern Ontario.

807. Crowe KA, Parker WH. 2008. Using portfolio 

theory to guide reforestation and restoration under climate change 

scenarios. Climatic Change 89:355-370.

Type: Journal 

Geographic Area: Central Canada

Compilers’ Keywords: decision support system, adaptation, 

Picea glauca, uncertainty, focal point seed zone, genotype, natural migration, management planning, seed transfer guidelines

Abstract: The general problem addressed by this study is that 

of designing a decision support system for planned adaptation 

to climate change that uses the principles of modern portfolio 

theory to minimise risk and maximise return of adaptive actions in an environment of deep uncertainty over future climate 

scenarios. Here we show how modern portfolio theory can use 

the results of a climate change impact model to select an optimal set of seed sources to be used in regenerating forests of 

white spruce in an environment of multiple, equally plausible 

futures climates. This study shows that components of solutions are not selected to perform equally well across all plausible futures; but rather, that components are selected to specialize in particular climate scenarios. The innovations of this 

fragmentation and climate change. Here, we suggest amendments to existing strategies combined with a review of alternative seed-sourcing strategies that propose to mitigate against 

these drivers. We present a provenancing selection guide based 

on confidence surrounding climate change distribution modelling and data on population genetic and/ or environmental 

differences between populations. Revegetation practices will 

benefit from greater integration of current scientific developments and establishment of more long-term experiments is key 

to improving the long-term success. The rapid growth in carbon and biodiversity markets creates a favourable economic 

climate to achieve these outcomes.

Link: http://s3.amazonaws.com/academia.edu.documents/30181053/Breed_et_al_provenance_2012.pdf?AW

SAccessKeyId=AKIAIR6FSIMDFXPEERSA&Expires=1-

382558363&Signature=pGq3eRDG2LawgufWthZN6Z8H3L

w%3D&response-content-disposition=inline

805. Broadhurst LM, Lowe A, Coates DJ, 

Cunningham SA, McDonald M, Vesk PA, Yates 

C. 2008. Seed supply for broadscale restoration: maximizing 

evolutionary potential. Evolutionary Applications 1:587-597

Type: Journal 

Geographic Area: Australia

Keywords: evolutionary potential, genetic diversity, germplasm, provenance, restoration, seed quality

Abstract: Restoring degraded land to combat environmental 

degradation requires the collection of vast quantities of germplasm (seed). Sourcing this material raises questions related to 

provenance selection, seed quality and harvest sustainability. 

Restoration guidelines strongly recommend using local sources 

to maximize local adaptation and prevent outbreeding depression, but in highly modified landscapes this restricts collection 

to small remnants where limited, poor quality seed is available, 

and where harvesting impacts may be high. We review three 

principles guiding the sourcing of restoration germplasm: (i) 

the appropriateness of using ‘local’ seed, (ii) sample sizes and 

population characteristics required to capture sufficient genetic 

diversity to establish self-sustaining populations and (iii) the 

impact of over harvesting source populations. We review these 

topics by examining current collection guidelines and the evidence supporting these, then we consider if the guidelines can 

be improved and the consequences of not doing so. We find that 

the emphasis on local seed sourcing will, in many cases, lead 

to poor restoration outcomes, particularly at broad geographic 

scales. We suggest that seed sourcing should concentrate less 

on local collection and more on capturing high quality and 

genetically diverse seed to maximize the adaptive potential of 

restoration efforts to current and future environmental change.

Link: http://pages.uoregon.edu/ecolrest/Spring%20

2011%20Restoration%20Readings/Summer%20Course%20

Docs/BR_2008_restoration_seed_selection.pdf

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284 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

among selected parents, and quantitative genetics analyses 

to identify elite genotypes for further genetic improvement 

or the establishment of production populations. BWB utility 

is demonstrated using a retrospective study of Douglas-fir 

(Pseudotsuga menziesii) progeny tests consisting of offspring 

produced from 150 controlled crosses among 60 parents and 

established over three sites. The empirical results are supported by theoretical expectations demonstrating anticipated 

minimum genetic response compared with conventional 

approaches. The method’s simplicity offers an exceptional 

opportunity for the development of comparable breeding 

efforts in developing countries, advanced and new breeding programmes, and economically important and ‘minor’ 

species.

Link: http://www.ncbi.nlm.nih.gov/pubmed/19393127

810. Erickson VJ. 2008. Developing native plant 

germplasm for national forests and grasslands in the Pacific 

Northwest. Native Plants Journal 9:255-266.

Type: Journal 

Geographic Area: Pacific Northwest USA

Keywords: grassland restoration, prairie, herbicide, restoration 

methods, seed addition, native diversity, invasive species

Abstract: Revegetation needs on national forests and grasslands in the Pacific Northwest are extensive and are expected to increase in the future as a result of growing challenges 

such as climate change, severe wildfires, invasive plants, 

and at-risk species. In response to these needs, Region 6 of 

the USDA Forest Service has implemented a new program 

to develop native plant materials for restoration activities 

that will be economical, effective, and consistent with Forest 

Service policy to protect biological diversity and plant genetic resources. The program has required strategic investments 

and commitments to research, infrastructure, equipment, and 

technology transfer. This article summarizes methods for 

species selection, plant material needs projections, propagule collection, nursery culture, and agricultural seed production. Although many challenges and constraints remain, 

considerable success has been achieved due to a dedicated 

workforce and the strong nursery system and agricultural 

infrastructure in the Pacific Northwest. Plant materials are 

becoming increasingly available and affordable as nurseries 

and seed producers gain experience and knowledge in their 

culture and management. In addition, this program is benefiting rural economies and contributing to the development 

of secondary, non-federal markets for native plant material 

services and products.

[Note: Erickson VJ. 2009. Developing native plant germplasm for national forests and grasslands in the Pacific Northwest 

- corrected table 1. Native Plants Journal 10:302.]

research rests in demonstrating that the powerful and widely 

used principles of quantifying and planning for risk and return 

in the uncertain environment of asset markets can be applied 

successfully to serve the objectives of planned adaptation to 

climate change. 

Link: http://flash.lakeheadu.ca/~whparker/ClimaticChange2008reprint.pdf

808. Echt CS. 1997. Use of molecular genetic markers in forest management. In: Communicating the Role of 

Silviculture in Managing the National Forests: Proceedings 

of the National Silviculture Workshop. Randor (PA): USDA 

Forest Service, Northeastern Forest Experiment Station. 

General Techinical Report NE-238. p 134-139. 

Type: Government Document

Geographic Area: North America

Compilers’ Keywords: biodiversity, silviculture, sustainability, white pine, timber management

Abstract: When managing forests for biodiversity or sustainability, attention must be given to how silvicultural practices affect genetic diversity. A new generation of DNA based markers 

affords a greater detail of genetic analysis than previously possible. These new markers, SSRs or microsatellites, have been 

used to demonstrate genetic diversity and infer evolutionary 

history of red pine, something that has not been possible with 

other markers. SSR markers developed by the Forest Service 

Research Biotechnology Unit are also being used to monitor 

how methods of sustainable timber management affect genetic 

diversity and breeding patterns within white pine stands on the 

Menominee Indian reservation.

Link: http://www.treesearch.fs.fed.us/pubs/15168

809. El-Kassaby YA, Lstiburek M. 2009. Breeding without breeding. Genetics Research 91:111-120.

Type: Journal 

Geographic Area: Global

Compilers’ Keywords: algorithms, computer simulation, genotype, genetic models, phenotype, plant genetics, Pseudotsuga 

menziesii, genotype selection, framework

Summary: An innovative approach to tree breeding called 

‘breeding without breeding’ (BWB) is presented. The method, as applied on the material in hand, allows the capture of 

75-85% of the genetic response to selection attained through 

conventional programmes without the need to do any controlled pollination and simplified or possibly no experimental 

field testing: both considered to be the most resource-demanding activities in breeding programmes. BWB combines 

the use of genotypic or phenotypic pre-selection of superior 

individuals, informative DNA markers for fingerprinting 

and pedigree reconstruction of offspring to assemble naturally created full- and half-sib families resulting from mating 

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the summer season. Using standard seed zone deployment 

practices in conjunction with these scientific findings provided 

a more comprehensive approach for defining seed transfer 

guidelines of Sitka and interior spruce populations. This program provided an example of how the field of biotechnology 

was used to assist in defining a seed deployment strategy for a 

region where introgression is a common occurrence between 

Sitka and interior spruce populations. 

813. Hamann A, Gylander T, Chen P. 2011.

Developing seed zones and transfer guidelines with multivariate 

regression trees. Tree Genetics and Genomes 7:399-408.

Type: Journal 

Geographic Area: North America

Keywords: genetic diversity, tree improvement, ecological genetics, genecology, aspen, red alder

Abstract: Managing seed movement is an important component of forest resource management to minimize maladaptation of planting stock in forest plantations. Here, we describe a 

new approach to analyze geographic patterns of adaptive and 

neutral genetic variation in forest trees and to link this genetic 

information to geographic variables for the delineation of seed 

zones and the development of seed transfer guidelines. We apply multivariate regression trees to partition genetic variation, 

using a set of environmental or geographic predictor variables 

as partitioning criteria in a series of dichotomous splits of the 

genetic dataset. The method can be applied to any type of 

genetic data (growth, adaptive, or marker traits) and can simultaneously evaluate multiple traits observed over several 

environments. The predictor variables can be categorical (e.g., 

ecosystem of seed source), continuous (e.g., geographic or 

climate variables), or a combination of both. Different sets of 

predictor variables can be used for different purposes: In two 

case studies for aspen and red alder, we show (1) how latitude, 

longitude, and elevation of seed sources in a provenance trial 

can be used to develop simple seed transfer guidelines; (2) how 

ecosystem classes and elevation as predictor variables can be 

used to delineate seed zones and breeding regions; and (3) how 

climate variables as predictors can reveal adaptation of genotypes to the environments in which they occur. Partitioning of 

genetic variation appears very robust regarding the choice of 

predictor variables, and we find that the method is a powerful 

aid for interpreting complex genetic datasets.

Link: http://www.ualberta.ca/~ahamann/publications/pdfs/

Hamann_et_al_2011.pdf

814. Heinze B, Westcott R, Schmidt J, Glossl J. 

1996. Application of random amplified polymorphic DNA 

(RAPD) to detect genetic variation in Norway spruce. New Forests 

11:173-184.

Type: Journal 

Geographic Area: Europe

811. Fay MF. 2003. Using genetic data to help guide 

decisions about sampling. Chapter 5. In: Smith RD, Dickie 

JB, Linington SL, Pritchard HW, Probert RJ, editors. Seed 

Conservation: Turning Science into Practice. London, United 

Kingdom: Royal Botanical Gardens. p 89-96. 

Type: Book Section

Geographic Area: Global

Compilers’ Keywords: conservation, AFLP, genetic variation, 

seed bank, guidelines

Summary: Many new genetic fingerprinting techniques were 

developed in the 1990s, and some of these are now being increasingly used to study genetic variation in populations of 

wild plant species. Examples are presented of the use of AFLP 

(amplified fragment length polymorphism) and plastid microsatellites in the characterization of genetic variation in wild 

species. How this information can be used to inform sampling 

strategies for seed banking is discussed.

Link: http://www.kew.org/ucm/groups/public/documents/

document/ppcont_013767.pdf

812. Grossnickle SC, Sutton BCS, Fan S, King J. 

1997. Characterization of Sitka x interior spruce hybrids: a biotechnological approach to seedlot determination. The Forestry 

Chronicle 73:357-362.

Type: Journal 

Geographic Area: Western Canada

Keywords: biotechnology, molecular genetics, ecophysiology, Sitka spruce, interior spruce, introgression zone, seedlot 

deployment

Abstract: The Nass Skeena Transition is an area located in 

British Columbia (BC), Canada, where a large introgression zone occurs between Sitka spruce (Picea sitchensis

(Bong.) Carr.) and interior spruce (white spruce, Picea glauca

(Moench) Voss x Engelmann spruce, Picea engelmannii Parry 

ex. Engelm.). When these species hybridize in this transition 

area, forestry programs face special problems in identifying 

seedlots that have the desired genetic makeup of source material collected from traditional seed zones. This paper presents an overview of a program that combined the disciplines 

of molecular genetics and ecophysiology to assist in defining 

the genetic makeup of seedlots based on DNA analysis results. 

Research first identified chloroplast, mitochondrial and nuclear 

DNA probes that were used to estimate species components 

for Sitka x interior spruce hybrid seedlots. Seedlings produced 

from Sitka x interior spruce hybrid seedlots were then characterized for ecophysiological parameters of drought tolerance, 

freezing tolerance and gas exchange patterns. DNA analysis 

results were then compared to ecophysiological parameters 

and findings indicated that seed source DNA profiles were related to drought tolerance throughout the year, freezing tolerance during fall acclimation and gas exchange patterns during 

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breeding. New genes may be infused into breeding populations 

from material selected in natural forest reserves to increase diversity. Production plantations will be genetically more diverse 

than naturally regenerated stands.

816. Kung FH, Rink G. 1993.A graphic technique for identifying superior seed sources for central hardwoods. In: Gillespie 

AR, Parker GR, Pope PE, Rink G, editors. Ninth Central 

Hardwood Forest Conference. St. Paul (MN): USDA Forest 

Service, North Central Forest Experiment Station. General 

Technical Report NC-161. p 101-106.

Type: Government Document

Geographic Area: Central USA

Compilers’ Keywords: forest production, genotype selection, 

tool, plantation, seed zones

Introduction: To maximize forest production, foresters need 

to plant the best genotypes provided by forest geneticists. 

Where should the forest geneticist search for the best seed 

sources? How far can one go south, or north to find them? 

The answer may rely on the species and the location of the 

test plantation. For example, when black walnut trees were 

tested in Illinois, Indiana, Missouri, and southern Michigan, 

trees from southern seed sources grew faster than those of local 

and northern seed sources. However, in Iowa and Minnesota, 

trees from northern sources were generally larger than trees 

from southern sources, and in Kansas and Ohio, tree size was 

not associated with latitude of seed source (Bey 1973, 1979). 

As the number of plantation increases, we may find that the 

most southern plantation will have only northern seed sources 

and the most northern plantation will have only southern seed 

sources, we may not be able to make a simple statement as to 

the best seed zone for each plantation location. Since a picture 

is better than a thousand words, we suggest a simple graphic 

solution useful to answer these questions. 

Link: http://www.treesearch.fs.fed.us/pubs/15308

817. Lindgren D, Ying CC. 2000. A model integrating 

seed source adaptation and seed use. New Forests 20:87-104.

Type: Journal 

Geographic Area: Sweden, Europe

Keywords: adaptability, seed zone, provenance, seed orchard, 

Scots pine

Abstract: A conceptual model that considers the performance 

(adaptability) of a seed source (= an origin) and the location or 

range of its deployment is developed employing the Cauchy 

function. The model assumes that there exists an optimal site 

type for each provenance origin (genetic material), and that 

loss in performance is a function of the “distance” (a measure 

of increasing maladaptation) from the optimal site. The model 

requires the estimate of three parameters: a site requirement 

value that measures site type in one dimension; a measure of 

Keywords: Picea abies, genetic markers, clone identification, 

tissue culture, segregation

Abstract: The RAPD assay is a screening method for genetic 

variation; in Norway spruce biology research, it may find 

many applications. We have investigated into its suitability for 

Norway spruce genetics by optimizing the protocol, testing 

for stability in clones, especially tissue culture clones, assessing the variation in a seed sample, and checking for correct 

Mendelian segregation in haploid megagametophytes. The 

RAPD assay produced numerous genetic markers quickly. The 

data obtained gave insight into the genetic make-up of clones 

and seed.

815. Kitzmiller JH. 1990. Managing genetic diversity in a 

tree improvement program. Forest Ecology and Management 

35:131-149.

Type: Journal 

Geographic Area: Southwestern USA

Compilers’ Keywords: reforestation, seed collection, silviculture, seed production

Abstract: Maintenance of species and within-species diversity is a goal of the USDA Forest Service, Pacific Southwest 

Region, which is examining and monitoring the genetic effects 

of silvicultural activities using isozyme analysis in conjunction with traditional methods. An effective monitoring and 

evaluation program will help improve silvicultural practices. 

Regional policies on gene conservation focus mainly on artificial regeneration. Tree improvement, nursery, and reforestation activities in the PSW Region may significantly affect 

adaptability, genetic diversity, and economic timber traits. 

The Base-Level Tree Improvement Program (TIP) aims for: 

(1) high adaptability via use of native species and local wild 

seed; (2) high within-species genetic diversity via designs for 

sampling several local seed stands and via long term plans for 

improvement using simple recurrent selection and bulking 

among the seed-production-area plantations; and (3) moderate 

improvement in performance via mild selection for superior 

phenotypes. The ceiling on genetic diversity is established by 

the seed collected. Nursery and reforestation practices may result in some loss of original (seed collection) diversity. Nursery 

practices aim to maximize plantable seedlings per unit of seed 

and thereby preserve diversity through sound management. 

Reforestation practices aim to achieve high local diversity by 

mixing appropriate species at planting, allowing natural regeneration to established in plantations, encouraging high survival 

and using higher planting densities so that natural selection 

can operate effectively. The High-Level TIP involving selective breeding and orchards will supplant the wild collections 

for major species and productive sites. Orchard seed will have 

a base of 50-60 unrelated, interbreeding parents, each representing a separate stand. Cooperative programs will enhance 

the genetic base by providing a broad range of material for 

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longitude) accounted for nearly half the variation in site index. 

Path analyses estimated a strong correlation (0.76) between 

genotype and environment, reflecting the steep adaptive clines 

that are well known for this species. The analysis also indicated 

that the genotype is about a third more important than the contemporary environment in determining phenotypic variation in 

dominant height of natural stands. The results suggest that a 

combined genetic/mensuration model has potential to improve 

both tree breeding efforts and stand management.

820. Neale DB, Wheeler NC. 2004. Mapping of quantitative trait loci in loblolly pine and Douglas-fir: a summary. Forest 

Genetics 11:173-178.

Type: Journal 

Geographic Area: USA

Keywords: QTLs, genomics, quantitative traits

Abstract: Classical quantitative genetic approaches have 

yielded a depth of understanding of the heritability and partitioning of variance of important traits in forest trees. Such 

knowledge has been applied in the genetic improvement of 

many species. Genetic improvement based on phenotypic selection in forest trees is rather slow however due to the long 

generation time in trees and the time to phenotypic evaluation. 

Genetic marker based approaches to tree improvement would 

increase the speed and precision of breeding. For this goal to 

be realized, the genes controlling quantitative traits must be 

identified. We summarize our progress toward achieving this 

goal based on a series of quantitative trait locus mapping experiments in two important conifer species. The path toward 

discovery of economically important alleles is now clear.

Link: http://www.tuzvo.sk/files/fg/volumes/2004/FG11-

34_173-178.pdf

821. O’Neill GA, Aitken SN. 2004. Area-based breeding zones to minimize maladaptation. Canadian Journal of Forest 

Research 34:695-704.

Type: Journal 

Geographic Area: Western Canada

Compilers’ Keywords: white spruce, Engelmann spruce, 

Picea glauca, Picea engelmannii, GIS, genetic adaptation, delineation method

Abstract: A new breeding zone delineation scheme identifies 

for a given number of zones the zone-boundary placement that 

minimizes regional maladaptation in breeding programs. First, 

an adaptive map is created by using conventional genetic test 

data. Then, the large array of predicted adaptive values is subjected to cluster analysis, which assigns each grid cell of the 

region to one of a predetermined number of clusters (breeding 

zones) such that the sum of the squared distances between each 

cell’s adaptive value and its cluster mean is minimized. This 

approach minimizes the average adaptive distance between 

optimal performance; and a flexibility measure of the width 

of seed source adaptability. The Cauchy function has a known 

integral, thus the average adaptability over a range (a possible 

seed use zone) can be mathematically evaluated. The model 

was also extended to seed orchard crops representing progeny of parents of variable origins. Scots pine information in 

Sweden was used to demonstrate possible applications of the 

model.

818. Mangold RD, Libby WJ. 1978. A model for reforestation with optimal and suboptimal tree populations.

Silvae Genetica 27:66-68.

Type: Journal 

Geographic Area: Global

Keywords: reforestation model, developmental growth strategy, optimal and suboptimal populations, nonlocal seed

Summary: A hypothetical reforestation model is presented 

which postulates growth rates by “optimal” and “suboptimal” 

tree populations. Populations of trees from the central zone of 

a species’ ecological tolerance may possess inherently greater 

capacities for rapid growth due to a “non-conservative” developmental growth strategy. Such a strategy probably evolves 

where environmental conditions are rarely limiting for growth, 

but biotic (competitive) interactions between neighboring trees 

are intense. When these populations are planted in less favorable sites, they may outgrow native populations. A method for 

determining the extent of planting zones for such populations 

is also presented. 

Link: http://www.silvaegenetica.de/fileadmin/content/dokument/archiv/silvaegenetica/27_1978/27-2-66.pdf

819. Monserud RA, Rehfeldt GE. 1990. Genetic and 

environmental components of variation of site index in inland 

Douglas-fir. Forest Science 36:1-9.

Type: Journal 

Geographic Area: Northern USA

Keywords: path analysis, provenance

Abstract: Results from two disparate and independent studies of inland Douglas-fir (Pseudotsuga menziesii [Mirb] Franco 

var. glauca) were combined to answer the following questions: 

First, how much variation in site index is associated with genetic variation? Second, what is the relative importance of the 

genetic and environmental components of phenotypic variation in explaining differences in mean height among stands of 

mature trees? Third, what is the magnitude of the interaction? 

An index to genetic variation, based on 3-year seedling height 

in provenance tests, accounted for approximately 40% of the 

variation in both 50-year and 100 year dominant height (i.e., 

site index) among 135 natural stands in northern Idaho and 

western Montana. Combining the genetic index and simple 

environmental variables (elevation, habitat series, latitude, 

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the viability of conserved populations, but requires the forfeiture of significant revenue from timber concessions.

Link: http://www.researchgate.net/publication/226767388_

Strategies_for_genetic_conservation_of_trees_in_the_Peruvian_Amazon/file/60b7d5147da1149c2e.pdf

823. Parker WH. 2000. Rates of change of adaptive variation in Picea mariana visualized by GIS using a differential systematic coefficient. New Forests 20:259-276.

Type: Journal 

Geographic Area: Ontario, Canada

Keywords: black spruce, breeding zones, Differential 

Systematic Coefficient, genecology, provenance test, delineation method

Abstract: Data from a short-term provenance trial of black 

spruce (Picea mariana (Mill.) B.S.P.) were used to illustrate 

a methodology to help locate breeding zone boundaries using 

adaptive variation models. Four height growth and survival 

variables were summarized by principal components analysis 

(PCA), and the first three axis scores were regressed against 

climate data determined from a recently developed Ontario 

Climate Model. The regression equations were used to model 

the PCA axes, and these models were interpreted as the three 

main components of adaptive variation in the data. These models were converted to geographic grids using GIS software. In 

a manner similar to that proposed for differential systematics 

applications, the Differential Systematic Coefficient (DSC) 

was adapted to be an indicator of the weighted average rate of 

change of clinally expressed adaptive variation over distance. 

An output grid was determined based on the DSC values, such 

that grid cells with higher coefficient values were made to appear darker on the resultant map; thus, the shaded areas corresponded to steeper portions of the clines of adaptive variation 

and serve as desirable indicators of the best locations for breeding zone boundaries.

824. Raymond CA, Lindgren D. 1990. Genetic 

flexibility: a model for determining the range of suitable environments for a seed source. Silvae Genetica 39:112-120.

Type: Journal 

Geographic Area: USA, Sweden

Keywords: breeding zones, Pinus sylvestris, genotype by environment interaction

Abstract: A mathematical model describing the relative performance of different seed sources over a range of environments has been developed and tested using data from Scots 

pine in Sweden and white ash in USA. The components of the 

model have a direct biological interpretation in terms of the 

fundamental nature of the problem: i.e., the maximal performance of a seed source, where this maximum will occur and 

the range of suitable sites for such a seed source. Comparisons 

the origin of a breeding program’s selected trees and planting 

locations throughout the region of focus. The procedure is illustrated by the use of adaptive values of 69 interior spruce 

(Picea engelmannii Parry ex Engelm. × Picea glauca (Moench) 

Voss) open-pollinated families (sources) from southeast 

British Columbia, Canada. Adaptive values of each 1.5 km × 

1.5 km grid cell in the 80,000-km2 region were predicted using 

a genecological model (R2 = 0.64), and the values were subjected to cluster analysis to identify breeding zone boundaries 

that were then mapped using a geographic information system. 

Regardless of the number of zones created, a regional maladaptation index was consistently smaller when zones were devised 

with area based cluster (ABC) analysis than when zones were 

created by dividing the region into bands of equal elevational 

or adaptive-value widths. Application of the ABC procedure 

should assist in identifying the optimum breeding-zone alignment for a given number of zones.

822. O’Neill GA, Dawson I, Sotelo-Montes C, 

Guarino L, Guariguata M, Current D, Weber JC. 

2001. Strategies for genetic conservation of trees in the Peruvian 

Amazon. Biodiversity and Conservation 10:837-850.

Type: Journal 

Geographic Area: Peru, South America

Keywords: agroforestry, genetic conservation, genetic erosion, 

Peruvian Amazon, tree genetic resources

Abstract: Forestry practices and high rates of land clearance for agriculture are causing genetic erosion of valuable 

tree species in the Peruvian Amazon, thereby endangering 

the economic sustainability of rural communities and limiting 

Peru’s opportunities for the development of new timber and 

non-timber forest products. The potential utility and limitations of six low-input interventions to help forestall further genetic erosion in the region are discussed, with a focus on local 

community involvement. Improved agroforestry systems may 

help reduce deforestation by increasing farm productivity, although methods to increase the currently low adoption rate of 

these technologies need to be developed. Use of strategic tree 

domestication techniques can also improve farm productivity 

and prevent inadvertent genetic drift and inbreeding associated with traditional domestication practices, although to have 

a major impact, current programs need to be extended across 

the region. Woodlot forestry could supplant selective extraction of timber and offers an attractive opportunity for poverty 

alleviation if appropriate credit and land tenure policies can be 

developed. However, it may also result in increased deforestation if activities on public land cannot be controlled. The implementation of improved seed collection systems and simple 

seed transfer guidelines would help to reduce the collection 

of seed of poor quality and low genetic diversity, and avoid 

maladapted plantings, although such programs are difficult to 

monitor and seed costs may increase. Strategic identification 

and design of in situ conservation areas would help to ensure 

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landscaping, and bioremediation requires knowledge of the 

adaptation of each species and, more specifically, knowledge 

of the adaptation of cultivars, strains, accessions, or ecotypes 

of a species to specific sites or regions. For agronomic and horticultural plants, specific adaptation information has been and 

continues to be developed by extensive testing. Rangeland, 

grassland, park, and restoration project managers often lack 

the resources to determine adaptation areas for plant materials because of the large number of species that are used and 

the extensive geographical areas that are serviced. Problems 

often arise in delineating adaptation areas for plant materials of 

both native and introduced species. Since ecoregion and plant 

hardiness zone classification systems integrate many climatic 

and geographic variables that determine plant adaptation, 

these 2 systems can be combined to develop Plant Adaptation 

Regions (PARs). A PAR map of the contiguous United States 

was developed by merging a widely used ecoregion map with 

the USDA Plant Hardiness Zone map, and is available in GIS 

format. Based on their geographic origins and/or test results, 

plant materials and their general areas of adaptation can be 

classified using PARs.

Link: http://www.researchgate.net/publication/43257641_

Plant_Adaptation_Regions_Ecological_and_Climatic_Classification_of_Plant_Materials/file/79e41512b945adf415.pdf

827. Wright JW. 1978. A simplified design for combined 

provenance and progeny testing. Silvae Genetica 27:68-70.

Type: Journal 

Geographic Area: Global

Keywords: provenance test, design

Summary: It is often desirable to combine the functions of 

provenance and progeny tests into one experiment testing 

the offspring of several different trees from each of many 

stands. A design is proposed whereby such an experiment 

may be simplified. The proposal is a variation of the compact-family design, with plots representing stands randomized within blocks, small sub-plots representing families 

within stands, and sub-plots systematically arranged within 

plots. The systematic arrangement is obtained by color coding when planting. That feature permits simplifications in 

the labeling, mapping and record keeping procedures without a sacrifice in quality of data. With the proposed design, a 

combined provenance-progeny test can be conducted almost 

as simply as an ordinary provenance test.

can thus be made between seed sources as to their site preferences and the relative flexibility of each seed source calculated. 

Decisions regarding the optimal deployment of seed may thus 

be simplified and such information used to refine the allocation 

of breeding zones. The model requires that a one-dimensional 

environmental gradient be defined. Such gradients may relate 

to a single overriding environmental factor (e.g. winter cold) 

which can be described by geographical information such 

as latitude, altitude and distance from the sea. Definition of 

this gradient is a necessary prerequisite to application of this 

model. Methods for establishing such gradients are discussed. 

Link: http://www.silvaegenetica.com/fileadmin/content/dokument/archiv/silvaegenetica/39_1990/39-3-4-112.pdf

825. St. Clair JB, Johnson GR, Erickson VJ, Johnson 

RC, Shaw NL. 2007. Seed zones for maintaining adapted 

plant populations. In: Norcini JG, editor. Native Wildflower 

Seed Production Research Symposium. Quincy (FL): University 

of Florida, Institute of Food and Agriculture, North Florida 

Research & Education Center. p 1.

Type: Conference Paper

Geographic Area: Western North America

Compilers’ Keywords: seed transfer guideline, native plants, 

genetics, common garden

Abstract: Seed zones delineate areas within which plant 

materials can be transferred with little risk that they will be 

poorly adapted to their new location. They ensure successful 

restoration and revegetation, and help maintain the integrity of 

natural genetic structure. The value of seed zones is recognized 

in numerous policy statements from federal and state agencies. Results from common garden studies indicate that local 

sources are often best adapted to local environments, although 

the degree of local adaptation differs between species and between traits within species. Seed zones have been used for over 

60 years in forest trees, whereas seed zones for grasses, forbs 

and shrubs used in restoration are largely lacking. We discuss 

several studies that have been completed or initiated to study 

adaptive genetic variation and the development of seed zones 

in species widely used in restoration in the western United 

States.

Link: http://www.treesearch.fs.fed.us/pubs/33640

826. Vogel KP, Schmer MR, Mitchell RB. 2005. Plant 

adaptation regions: ecological and climatic classification of plant 

materials. Rangeland Ecology and Management 58:315-319.

Type: Journal 

Geographic Area: USA

Keywords: ecoregion, plant hardiness zone, ecotypes, plant 

germplasm, cultivars, native plants

Abstract: The effective use of plant materials for an array 

of objectives including conservation, restoration, renovation, 

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828. Alden JN. 1991. Provisional tree seed zones and transfer guidelines for Alaska. Portland (OR): USDA Forest Service, 

Pacific Northwest Research Station. General Technical Report 

PNW-GTR-270. 35 p.

Type: Government Document

Geographic Area: Alaska, USA

Keywords: Alaska, seed zone, seed transfer and labeling guidelines, tree distribution, tree limit, forest area, genetic diversity, 

climate, physiography

Abstract: Four hundred and eighty-six provisional tree seed 

zones were delineated within 24 physiographic and climatic 

regions of Alaska and western Yukon Territory. Estimated forest and potential forest land within altitudinal limits of tree species in Alaska was 51,853,000 hectares (128,130,000 acres). 

Seed transfer guidelines and standard labeling of seed collections are recommended to reduce losses from maladaptation 

and maintain the genetic identity and productivity of future 

forests in Alaska.

Link: http://www.treesearch.fs.fed.us/pubs/25389

829. Atlas of Living Australia. (URL accessed 07 

November 2015) 

Type: Web Page

Geographic Area: Australia

Compilers’ Keywords: climate change, restoration, seed 

source, native plants, decision support system

Synopsis: The Atlas of Living Australia (Atlas) contains information on all the known species in Australia aggregated 

from a wide range of data providers: museums, herbaria, community groups, government departments, individuals and universities. This site can be used to access information pages for 

each species containing photos, descriptions, maps and observations; access scientific and common names; explore the flora 

and fauna reported around your neighbourhood; learn about 

Australia’s biodiversity collections at museums, herbaria and 

other institutions; learn about citizen science projects, map, 

analyze and visualize biodiversity and environmental data and 

trends; access tools to help track changes in biodiversity and 

the environment; download and use open source tools; download biodiversity data; access images, literature and genetic 

information through Australian nodes of international data repositories; and volunteer for digitization projects. As well, you 

can get facts and figures about the information contained in the 

Atlas from the website dashboard. 

Link: http://www.ala.org.au/

General (583–616)—reviews, history, 

summaries, justification

Research (617–802)—genetic studies, tree 

improvement, climate change

Strategies (803–827)—framework, delineation, modelling

Resources (828–877)—guidelines, zones, 

maps, tools, software

Transfer Guidelines

and Zones

#. Author(s), Date. Title. Source.

Type: 

Geographic Area: 

Keywords: 

Description:

Link:

location of study

keywords written by the 

author(s) or provided by the 

compilers

ID number and

information

abstract, summary, or 

portions of the introduction 

or conclusion written by the 

author(s) or provided by the 

compilers

book, book section, conference 

paper, dissertation, government 

document, journal article, 

newsletter, presentation, report, 

thesis or web page

internet address of reference

Each reference contains the following:

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USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 291

successful restoration project. The key to identifying appropriate plant materials (e.g., seeds) lies in understanding the genetics of adaptation through common garden studies or reciprocal 

transplant experiments. However, restoration practitioners are 

frequently required to make seed collection and deployment 

decisions for species and landscapes which lack seed zones 

and transfer guidelines established through genetic research. 

So what are practitioners to do when no seed transfer guidelines exist for a species of interest? One approach for choosing 

germplasm in the absence of genetic information is to try and 

match the seed source and planting site conditions as closely 

as possible. Our provisional seed zone model provides a powerful decision support tool for characterizing site conditions 

and choosing plant material sources that originate from similar 

environments. The model integrates climate factors that are 

known to affect plant survival and growth (temperature and 

precipitation) and ecological factors. It builds on earlier efforts 

such as USDA plant hardiness zones (Cathy 1990, www.arborday.org) and “Plant Adaptation Regions” (PARs, Vogel et 

al. 2005). The provisional zones can be utilized to guide movement of plant material until species specific information becomes available via genetic research studies on adaptive traits.

Link: http://www.fs.fed.us/wwetac/threat_map/SeedZones_

Intro.html

833. Bower AD, St. Clair JB, Erickson VJ. 2010.

Generalized provisional seed zones for native plants. In: National 

Native Seed Conference: Native Plant Materials Development, 

Production and Use in Habitat Restoration. Corvallis (OR): 

Institute for Applied Ecology. 28 p.

Type: Presentation

Geographic Area: USA

Compilers’ Keywords: seed transfer zones, provisional zones, 

ecoregion, climate change, genetic information, seed transfer 

guideline, plant materials, adaptation 

Abstract: Deploying vigorous, well adapted, and ecologically appropriate plant materials is a core component of a successful restoration project. The key to identifying appropriate 

plant materials lies in understanding the genetics of adaptation 

through common garden studies. However, restoration practitioners often deploy plant species on the landscape for which no 

seed transfer guidelines have been established through genetic 

research. What are practitioners to do when no seed transfer 

guidelines have been established for a species of interest? We 

have developed generalized provisional seed zones that can be 

applied to any plant to help guide seed movement. These seed 

zones area based on the intersection of high resolution (800 m 

x 800 m cell size) observed climatic data. The intersection of 

winter minimum temperature and annual precipitation delineates zones for trees, shrubs, and woody plants, while the intersection of average maximum termperature and precipitation is 

used for zones for grasses and herbaceous plants. The resulting 

830. Beaulieu J. 2009.Optisource: a tool for optimizing seed 

transfer. Quebec, Canada: Natural Resources Canada, 

Canadian Forest Service 55. 2 p.

Type: Government Document

Geographic Area: Eastern Canada 

Compilers’ Keywords: seed production, tool

Introduction: The ministère des Ressources naturelles et de 

la Faune du Québec (MRNF) oversees the production of tree 

seed and seedlings to meet the reforestation needs of public and 

private forests. Nearly 600 million viable seeds are required 

for forest regeneration every year. Tree seeds are produced in 

a variety of locations in Quebec. Optisource, the decision support tool used by the MRNF, was designed by Canadian Forest 

Service researchers using a geographic information system. This 

software combines data concerning ecological districts with data 

obtained from seed source transfer models. When improved or 

local seed sources are not available to reforest a given site and 

a different source needs to be used, Optisource can be used to 

determine which seed sources are best suited to the planting site.

Link: http://cfs.nrcan.gc.ca/pubwarehouse/pdfs/31544.pdf

831. Bower AD, St. Clair JB, Erickson VJ. 2008.

Delineation of provisional seed zones for native plants.

Portland (OR): USDA Forest Service. 

Type: Map/Government Document

Geographic Area: USA

Compilers’ Keywords: provisional seed zones, native plants, 

restoration, seed transfer guideline

Introduction: Deploying vigorous, well adapted, and ecologically appropriate plant materials is a core component of a 

successful restoration project. The key to identifying appropriate plant materials (e.g., seeds) lies in understanding the genetics of adaptation through common garden studies. However, 

restoration practitioners often deploy plant species on the landscape for which no seed transfer guidelines have been established through genetic research. So what are practitioners to do 

when no seed transfer guidelines have been established for a 

species of interest?

Link: http://www.fs.fed.us/wildflowers/nativeplantmaterials/rightmaterials.shtml

832. Bower AD, St. Clair JB, Erickson VJ. 2008.

Provisional seed zones for native plants. USDA Forest Service. 

5 p.

Type: Government Document

Geographic Area: USA

Compilers’ Keywords: provisional seed zones, native plants, 

restoration, seed transfer guidelines

Background: Deploying vigorous, well adapted, and ecologically appropriate plant materials is a core component of a 

CHAPTER 4 • TRANSFER GUIDELINES AND ZONES • RESOURCES
292 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Compilers’ Keywords: seed transfer guidelines, forest management, seed collection

Introduction: California forest tree seed zones were established originally by Fowells (1946), with revisions by Roy 

(1963) and Schubert (1966). The Forest Tree Seed Committee of 

the Northern California Section, Society of American Foresters, 

has revised the original zones and updated the recording systems 

described in the earlier reports. The Forest Tree Seed Committee 

acknowledges the assistance of foresters and tree seed dealers in 

California and the Pacific Northwest who reviewed the revised 

zone map and reporting system. The California Tree Seed Zone 

map is an integral part of the se3ed collection reporting system. 

Zones are delineated on the basis of collection criteria adopted 

in the USDA forest seed policy of 1939. The policy specifies 

that seed should 1) be collected within 100 miles north or south 

of the planting site and 2) differ in elevation by less than 1000 

feet. Also, 3) careful consideration should be given to areas having a unusual climatic, topographic, or soil conditions that might 

greatly affect tree growth. On the basis of these three criteria, the 

State was divided into six physiographic and climatic regions, 

32 subregions within regions, and 85 seed collection zones. 

One region is divided into four physiographic and climatic areas having 18 of the 85 zones. Seed collection zones are limited 

to about 50 miles in latitude. Where possible, boundaries have 

been chosen to follow natural features, such as crests of mountain ranges, ridgetops, and rivers, or physical features such as 

highways, canals, and railroads. The seed collection zones and 

three-digit method of designating the various zones are coordinated with the map and coding system developed by the Western 

Forest Tree Seed Council in 1966 and now used in Oregon and 

Washington.

Link: http://www.treesearch.fs.fed.us/pubs/41438

836. Burns KS, Schoettle AW, Jacobi WR, 

Mahalovich MF. 2007. White pine blister rust in the 

Rocky Mountain Region and options for management. Golden 

(CO): USDA Forest Service, Rocky Mountain Research Station. 

Biological Evaluation R2-07-04. 38 p.

Type: Government Document

Geographic Area: Western USA

Compilers’ Keywords: forest management, Pinus

Introduction: White pine blister rust (WPBR) is an exotic, 

invasive fungal disease of white, stone, and foxtail pines (also 

referred to as white pines or five-needle pines) in the genus 

Pinus and subgenus Strobus (Price et al. 1998). The disease, 

which is native to Asia, was accidentally introduced separately into eastern and western North America at the beginning 

of the 20th century. In the west, blister rust was introduced 

on infected eastern white pine (Pinus strobus) nursery stock 

shipped to Vancouver, B.C. from France in 1910. Since then, 

white pine blister rust has spread throughout the distributions of sugar pine (P. lambertiana), western white pine (P. 

seed zones represent areas of relative climatic similarity, and 

movement of seed within these zones should help to minimize 

maladaptation. Superimposing Omernik’s level III ecoregions 

over these seed zones helps to distinguish areas that are similar 

climatically yet different ecologically. These provisional seed 

zones should be considered a starting point as guidelines for 

seed transfer, and should be utilized in conjunction with appropriate species-specific information as well as local knowledge 

of microsite differences.

Link: http://www.fs.fed.us/rm/boise/research/shrub/projects/PowerPoint_Presentations/2011/bower.pdf

834. Bower AD, St. Clair JB, Erickson VJ. 2014.

Generalized provisional seed zones for native plants. Ecological 

Applications 24:913-919.

Type: Journal Article

Geographic Area: USA

Keywords: adaptive traits, aridity, ecoregion, genetic variation, 

local adaptation, native plants, precipitation, restoration, seed 

transfer guideline, seed zone, temperature

Abstract: Deploying well-adapted and ecologically appropriate plant materials is a core component of successful 

restoration projects. We have developed generalized provisional seed zones that can be applied to any plant species 

in the United States to help guide seed movement. These 

seed zones are based on the intersection of high-resolution 

climatic data for winter minimum temperature and aridity 

(as measured by annual heat : moisture index), each classified into discrete bands. This results in the delineation of 

64 provisional seed zones for the continental United States. 

These zones represent areas of relative climatic similarity, 

and movement of seed within these zones should help to 

minimize maladaptation. Superimposing Omernik’s level III 

ecoregions over these seed zones distinguishes areas that are 

similar climatically yet different ecologically. A quantitative 

comparison of provisional seed zones with level III ecoregions and provisional seed zones within ecoregions for three 

species showed that provisional seed zone within ecoregion 

often explained the greatest proportion of variation in a suite 

of traits potentially related to plant fitness. These provisional 

seed zones can be considered a starting point for guidelines 

for seed transfer, and should be utilized in conjunction with 

appropriate species-specific information as well as local 

knowledge of microsite differences.

Link: http://www.treesearch.fs.fed.us/pubs/47587

835. Buck JM, Adams RS, Cone J, Conkle MT, 

Libby WJ, Eden CJ, Knight MJ. 1970. California tree 

seed zones. San Francisco (CA): USDA Forest Service. 7 p.

Type: Government Document

Geographic Area: California, USA

CHAPTER 4 • TRANSFER GUIDELINES AND ZONES • RESOURCES
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 293

Abstract: Provisional seed zones and breeding zones were 

developed for white pine (Pinus monticola Dougl. ex D. Don) 

in the Cascade Range of western Washington and Oregon. 

Recommendations were based on genetic variation patterns 

obtained by evaluating genotypes of trees from 115 locations. 

Genotypes controlling growth vigor and growth rhythm were 

assessed in common gardens. Five zones, delimited mainly 

by latitude, were recommended: two in Washington, two in 

Oregon, and one in both States adjoining the Columbia River.

Link: http://www.treesearch.fs.fed.us/pubs/9240

839. Campbell RK, Sugano AI. 1993. Genetic variation and seed zones of Douglas-fir in the Siskiyou National Forest. 

Portland (OR): USDA Forest Service, Pacific Northwest Region. 

Research Paper PNW-RP-461. 29 p.

Type: Government Document

Geographic Area: Oregon, USA

Abstract: Pseudotsuga menziesii, geographic genetic variation, 

adaptation, ecological genetics, genecology

Abstract: Provisional seed zones and breeding zones were developed for Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) 

in the Siskiyou National Forest in southwestern Oregon. Zones 

were based on maps of genetic variation patterns obtained by 

evaluating genotypes of trees from 260 locations in the region. 

Genotypes controlling growth vigor and growth rhythm were 

assessed in a common garden. Within the Forest, three breeding blocks were recommended, with different numbers of elevational bands in each block: from 0 to 610 meters, from 611 

to 838 meters, and then a series of bands 152 meters wide at 

higher elevations.

Link: http://www.treesearch.fs.fed.us/pubs/20575

840. Cathey HM. 2012. USDA plant hardiness zone map. 

Washington (DC): USDA National Arboretum, Agricultural 

Research Service. 

Type: Map/Government Document

Geographic Area: USA

Compilers’ Keywords: plant hardiness zone, map, transfer 

guidelines, temperature delineation

Summary: The 2012 USDA Plant Hardiness Zone Map is the 

standard by which gardeners and growers can determine which 

plants are most likely to thrive at a location. The map is based 

on the average annual minimum winter temperature, divided 

into 10 °F zones.

Link: http://planthardiness.ars.usda.gov/PHZMWeb/

Downloads.aspx

841. Cunningham RA. 1975. Provisional tree and shrub 

seed zones for the Great Plains. Fort Collins (CO): USDA Forest 

Service, Rocky Mountain Forest and Range Experiment Station. 

Publication 71. 21 p.

monitcola) and eastern white pine and all but the southern 

extents of whitebark pine (P. albicaulis) and limber pine (P. 

flexilis) and the western extent of southwestern white pine 

(P. strobiformis). WPBR has not been found in most of the 

Rocky Mountain bristlecone pine (P. aristata) range and is yet 

to be found on Great Basin bristlecone pine (P. longaeva). It 

was once thought that the remote dry habitats occupied by 

these species would not support rust establishment, however 

WPBR can now be found at many of these sites. Cronartium 

ribicola, the fungus that causes WPBR, requires an alternate 

host, currants and gooseberries in the genus Ribes and possibly species of Pedicularis and Castilleja (McDonald et al. 

2006, Zambino et al. 2007) to complete its life cycle. WPBR 

infects Ribes seasonally causing minimal damage such as leaf 

spots and premature defoliation; the infections are shed each 

year with leaf abscission. The disease is perennial on infected 

pines causing cankers that usually lead to mortality. WPBR 

has caused widespread decline and mortality over millions of 

acres resulting in dramatic changes in successional pathways 

and ecosystem functions and the disease continues to spread 

and intensify wherever five-needle pines occur.

Link: http://www.treesearch.fs.fed.us/pubs/29450

837. Campbell RK, Sugano AI. 1987. Seed zones and 

breeding zones for sugar pine in southwestern Oregon. Portland 

(OR): USDA Forest Service, Pacific Northwest Region. 

Research Paper PNW-RP-383. 18 p.

Type: Government Document

Geographic Area: Oregon, USA

Keywords: seed zones, geographic variation, genetic variation, 

adaptation (plant), sugar pine, Pinus lambertiana

Abstract: Provisional seed zones and breeding zones were 

developed for sugar pine (Pinus lambertiana Dougl.) in southwestern Oregon. Zones are based on a map of genetic variation patterns obtained by evaluating genotypes of trees from 

142 locations in the region. Genotypes controlling growth 

vigor and growth rhythm were assessed in a common garden. 

Within southwestern Oregon, two zones are recommended 

for low elevations (< 740 m), two zones for middle elevations (> 740 and < 1172 m), and four zones for high elevations 

(> 1172 m).

Link: http://www.treesearch.fs.fed.us/pubs/9219

838. Campbell RK, Sugano AI. 1989. Seed zones and 

breeding zones for white pine in the Cascade range of Washington 

and Oregon. Portland (OR): USDA Forest Service, Pacific 

Northwest Region. Research Paper PNW-RP-407. 26 p.

Type: Government Document

Geographic Area: Pacific Northwest USA

Keywords: genetic variation, geographic variation, adaptation 

(plant), seed-transfer, Pinus monticola

CHAPTER 4 • TRANSFER GUIDELINES AND ZONES • RESOURCES
294 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Keywords: red alder, Alnus rubra, tree improvement, quantitative genetics, genecology, resource management

Abstract: During the last two decades, the value of red alder 

wood products has substantially increased and several initiatives have been launched in the United States to use red alder 

for reforestation. Nonetheless, red alder is a largely neglected 

resource in British Columbia. This review paper examines 

the reasons behind the under-utilization of red alder in British 

Columbia and investigates whether changes in red alder management practices could improve the value of the resource. 

Red alder’s potential for plantation forestry and genetic tree 

improvement are discussed, and possible breeding objectives 

were evaluated with consideration for the species biology, 

growth, product value and market demand. Seed transfer rules 

and the possibility of gains from selection are summarized in 

the light of new research results in genecology and quantitative 

genetics for red alder populations in British Columbia.

Link: http://www.ualberta.ca/~ahamann/publications/pdfs/

Hamann_2001.pdf

845. Howe GT, St. Clair JB, Beloin R. 2009. Seedlot 

selection tool. Corvallis (OR): Oregon State University and 

USDA Forest Service. (URL accessed 29 November 2015)

Type: Web Page

Geographic Area: North America

Compilers’ Keywords: seed transfer zone, model, GIS, reforestation, climate change, tools, seed transfer guideline, delineation method

Introduction: The Seedlot Selection Tool is a joint effort 

of the Department of Forest Ecosystems and Society at the 

Oregon State University College of Forestry and the U.S. 

Forest Service Pacific Northwest Research Station. Funding 

comes from the U.S. Forest Service. The seedlot selection 

tool (SST) is a GIS mapping program designed to help forest 

managers match seedlots with planting sites based on climatic 

information for the Northwest, Midwest, Central, and East in 

North America. The tool can be used to map current climates, 

or future climates based on selected climate change scenarios. 

Although it is tailored for matching seedlots and planting sites, 

it can be used by anyone interested in mapping present or future climates defined by temperature and precipitation.

Link: http://sst.forestry.oregonstate.edu/index.html

846. Howe GT, Jayawickrama K, Cherry M, Johnson 

GR, Wheeler NC. 2006. Breeding Douglas-fir. Plant 

Breeding Reviews 27:245-353.

Type: Journal 

Geographic Area: Western North America

Compilers’ Keywords: plant breeding, forest management, 

silviculture, genetics, Pseudotsuga menziesii, economic value

Type: Government Document

Geographic Area: Central USA

Compilers’ Keywords: seed transfer zones, tree and shrub 

zones, transfer guidelines, seed collection, seed source

Preface: Seed collection zones are subdivisions of land areas 

established to identify seed sources and to control the movement of seed and planting stock. Seed zones are needed for 

many species because of the genetic variation associated with 

their geographic distribution. Zone boundaries may be delineated from experimental data that identify genetic variation, or 

by analysis of the environmental factors that most likely acted 

as selective forces in creating such genotypic variation. 

Link: http://www.treesearch.fs.fed.us/pubs/24593

842. Forest Seedling Network. 2012. (URL accessed 

07 November 2015) 

Type: Web Page

Geographic Area: USA

Compilers’ Keywords: seed zones, seed selection, forest management, seed transfer guidelines

Compilers’ Synopsis: Interactive website connecting forest 

landowners with seedling providers and forest management 

services and contractors. The site includes seed zone maps for 

Washington, Oregon, California, Montana, and Idaho.

Link: http://www.forestseedlingnetwork.com/

843. British Columbia Ministry of Forests. 1994.

Seed and vegetative material guidebook. British Columbia (CA): 

British Columbia Ministry of Forests.

Type: Government Document

Geographic Area: Western Canada

Compilers’ Keywords: seed transfer guideline, forest management, forest practices code, guidelines

Introduction: This guidebook has been prepared to help 

forestry practitioners meet the requirements of the Forest 

Practices Code with respect to planning, collecting, registering 

and using seed and vegetative material. The guide is relevant 

to all forestry operations that use planting or direct seeding as 

a method of reforestation. The procedures, processes and options described are arranged in the same order as the business 

of reforestation—from planning to planting. 

Link: http://www.for.gov.bc.ca/tasb/legsregs/fpc/fpcguide/

veg/seedtoc.htm

844. Hamann A. 2001. Utilization and management of 

red alder genetic resources in British Columbia. The Forestry 

Chronicle 77:705-712.

Type: Journal 

Geographic Area: Western Canada

CHAPTER 4 • TRANSFER GUIDELINES AND ZONES • RESOURCES
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 295

should be given to ecological considerations. Different species 

and seed sources will be best at different planting sites, and 

the challenge is therefore to match species and seed sources to 

planting site. Well-adapted seed sources may ensure a reliable 

yield for farmers, while maladapted seed sources may result in 

loss—or even total failure. In the present ‘Tree Planting Zones’ 

planting sites with similar environmental conditions are grouped 

together into zones for which specific seed sources can be developed and thereby increase farmers’ planting success. The 

‘Tree Planting Zones’ can be recognised in the field by farmers 

and will be utilized where there is the greatest potential for seed 

demand. The distribution of vegetation types is the best indicator of growing conditions for trees and bushes. ‘Tree Planting 

Zones’ have been developed in the warmer ecological zones 

where most planting by farmers take place and consequently 

where most seed transfers are required. In the colder ecological 

zones transfer of seeds for the relatively limited planting should 

be avoided and seed should be collected at the planting site.

Link: http://curis.ku.dk/ws/files/35157079/Case_Study_1_

Nepal.pdf

849. Mahalovich MF, Hoff RJ. 2000. Whitebark pine operational cone collection instructions and seed transfer guidelines.

Nutcraker Notes 11:10-13.

Type: Newsletter

Geographic Area: Western USA

Compilers’ Keywords: Pinus albicaulis, restoration, silviculture, seed transfer zones

Preface: This issue of Nutcracker Notes contains articles on 

the importance of genetics in whitebark pine research and 

management. On pages 10 through 13, Mahalovich and Hoff 

outline seed transfer guidelines and seed collection instructions 

for whitebark pine that are blister-rust resistant. They include 

criteria for collecting cones such as tree condition, form, age, 

cone condition, and caches. Other information provided are 

how to distinguish whitebark from limber pine, how to protect 

cones from predation and specific instructions on what parent 

trees to collect from. From isozyme studies, this species has 

little genetic diversity, thus is considered a generalist species. 

Seeds can be transferred within mountain ranges listed, but 

not across ranges. The guidelines are provisional, because of 

zone boundaries, limited genetic information, and issues associated with seed viability in storage, the author suggests a 

wider transfer from seed source, with emphasis on using seed 

collected from blister rust resistant trees: 1° latitude ± 50 miles, 

1° longitude ± 50 miles and no restriction on elevation.

Link: https://www.google.com/url?sa=t&rct=j&q=&esr

c=s&source=web&cd=1&ved=0CCoQFjAA&url=http%

3A%2F%2Ffedgycc.org%2FNutcrackerNotesNo.11.doc.

doc&ei=0IdyUrv1EOuajAKJkoHAAw&usg=AFQjCNGL8q

A5uWuNH8jROjwPqRpMd0xeUQ&sig2=Ju89mEFn-lKwni

43NYYgzw&bvm=bv.55819444,d.cGE

Objective: In this paper, we describe the (1) key elements that 

distinguish tree breeding from other types of crop breeding, (2) 

differences between Douglas-fir breeding and other types of 

tree breeding, (3) environmental, biological, and sociological 

factors that make Douglas-fir breeding unique, (4) current state 

of Douglas-fir breeding, and (5) future of Douglas-fir breeding and research needs. Although Douglas-fir improvement is 

practiced throughout the world, we will focus on Douglas-fir 

breeding in western North America (i.e., California, Oregon, 

Washington, British Columbia, Idaho, and Montana), and the 

Pacific Northwest in particular - where most of the Douglas-fir 

breeding is occurring. 

Link: http://www.fsl.orst.edu/pnwtirc/Electronic%20version%20of%20some%20pubs/(2006)%20Howe%20et%20

al%20%20Breeding%20Douglas-fir.pdf

847. Institute for Applied Ecology. Native seed 

network. Corvallis (OR): Institute for Applied Ecology. 

(URL accessed 25 November 2015)

Type: Web Page

Geographic Area: USA

Compilers’ Keywords: seed selection, ecoregions, plant selection, native plant materials, seed directory

Mission: The Native Seed Network promotes the use of native plants in ways that support the ecological integrity of 

both natural and manipulated ecosystems. The Native Seed 

Network advocates using native plant materials from appropriate genetic sources to conserve biological diversity, and to 

maintain the adaptive capability of ecosystems, communities 

and plant populations. The Native Seed Network works with 

public agencies, educators and private groups to increase the 

availability of affordable native plant materials. The website 

is an interactive database of native plant and seed information 

and guidelines for restoration, native plant propagation, and 

native seed procurement by ecoregion for the United States. 

Link: http://www.nativeseednetwork.org/

848. Lilleso JPB, Dhakal LP, Shrestha TB, Nayaju 

RP, Shrestha R, Dahl Kjaer E. 2001. Tree planting 

zones in Nepal—an ecological approach based on vegetation 

types. Kathmandu, Nepal: Danida Forest Seed Centre, Tree 

Improvement and Silviculture Component. DFSC Case Study 

No. 1, TISC Tech Paper 103. 45 p.

Type: Government Document

Geographic Area: Nepal, Southern Asia

Compilers’ Keywords: seed transfer, tree planting zones, silviculture, seed source, adaptation, guidelines

Summary: A substantial number of species (especially fodder 

trees) are planted by farmers and it is likely that the demand 

for many species will increase in the future. Given the extraordinarily large ecological variation in Nepal, special attention 

CHAPTER 4 • TRANSFER GUIDELINES AND ZONES • RESOURCES
296 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

seed/stock transfer, and 7) urge that public and private sectors 

follow this policy in all forest, shelter-belt, and erosion-control 

projects. The policy defines local as seed/stock from an area 

with similar climate conditions, or as that collected within 100 

miles and 1000 feet in elevation of the planting site. 

852. McKenney DW. 2002. Seedwhere: a computer tool 

for tree planting and ecological restoration. Sault Ste. Marie, 

Ontario, Canada: Natural Resources Canada, Canadian Forest 

Service, Northern Forestry Centre. Bulletin No. 21. 2 p.

Type: Government Document

Geographic Area: Canada

Compilers’ Keywords: seed transfer guidelines, genetic adaptation, GIS, planting zones, forest management, nursery stock, 

forest regeneration, climate change, restoration, tool

Introduction: It is generally recognized that plants are genetically adapted to the prevailing climate. Planting stock is 

often moved across environmental gradients with little knowledge of the ecological risks. For long-lived species (e.g., forest trees) this introduces risks of increased mortality, lowered 

potential growth rates, and increased susceptibility to insects 

and diseases. Thus mistakes in seed transfer in forest regeneration programs can be costly. Seedwhere is a Geographic 

Information System (GIS) tool designed to assist in making 

nursery stock and seed transfer decisions for forest regeneration activities over environmental gradients. Given a suitable 

database, Seedwhere maps out the similarity of a chosen location to the rest of the region of interest. The location could be a 

potential seed collection or planting area.

Link: http://cfs.nrcan.gc.ca/pubwarehouse/pdfs/20952.pdf

853. McKenney DW, Mackey BG, Joyce D. 1999.

Seedwhere: a computer tool to support seed transfer and 

ecological restoration decisions. Environmental Modelling 

14:589-595.

Type: Journal 

Geographic Area: Global

Keywords: climate grids, GIS, forest management, Arcview, 

Gower metric

Abstract: It is generally recognized that plants are genetically adapted to prevailing climate. However, there are very 

few genecological studies that quantify these relationships. 

Planting stock is often moved across environmental gradients 

with little knowledge of the ecological risks. For long-lived 

species (e.g., forest trees) this introduces risks of increased 

mortality, lowered potential growth rates, and increased susceptibility to insects and diseases. Seedwhere is a computerbased tool to support decisions on moving plant material 

across environmental gradients. The Gower similarity metric 

has been invoked as an Avenue extension in the ARCVIEW 

Geographic Information System. The program can be used 

850. Mahalovich MF, McArthur ED. 2004. Sagebrush 

(Artemisia spp.): seed and plant transfer guidelines. Native Plants 

Journal 5:141-148.

Type: Journal 

Geographic Area: North America 

Keywords: seed zones, adaptive traits, elevation gradients, ecotypes, ploidy, hybridization

Abstract: Seed and plant transfer guidelines are offered for 

11 species of sagebrush (Artemisia arbuscula, A. bigelovii, A. 

cana, A. rothrockii, A. longiloba, A. argillosa, A. rigida, A. nova, 

A. tripartita, A. pygmaea, and A. tridentata). These species 

constitute the true sagebrushes of the subgenus Tridentatae of 

Artemisia. The geographic distribution of each species serves 

as the geographic boundary for the 11 seed zones, with the additional restriction that seeds should not be moved farther than 

483 km (300 mi) to their target planting site, and if less than 

483 km, not outside of their native distribution. For A. tridentata, seed transfer should ensure that subspecies are planted 

with respect to moisture and elevational gradients. For all other 

species, no additional transfer guidelines are proposed based 

on changes in elevation, but when local data suggest moisture 

gradients and ranges of elevation in excess of 458 m (1,500 

ft), conservative guidelines could further restrict seed transfer 

up 153 m (500 ft) in elevation, or down 305 m (1,000 ft) in 

elevation, from the origin collection area. Correctly applied, 

seed and plant transfer guidelines minimize the risk of planting maladapted stock, increasing the survival and reproductive 

success to achieve restoration, rehabilitation, reclamation, and 

wildlife habitat improvement objectives.

Link: http://npj.uwpress.org/content/5/2/141.full.pdf+html

851. McCall MA, Silcox FA, Myer DS, Wallace HA. 

1939. Forest policy of U.S. Department of Agriculture. Journal 

of Forestry 37:820-821.

Type: Journal 

Geographic Area: USA

Compilers’ Keywords: seed transfer guidelines, restoration, 

seed zones, local adaptation, local seed, forest management, 

federal policy

Compilers’ Abstract: This is a research note written by the 

USDA Chairman of the Seed Policy Committee, Chief of the 

Forest Service, Acting Chief of the Soil Conservation Service, 

and Secretary. It reports USDA Forest Service policy guidelines on tree and shrub planting with respect to location and 

objective. The seven guidelines are 1) use seed and stock of 

known origin, 2) provide proof of seed and stock origin, 3) details of the seed and stock including lot number, year, species, 

origin, and proof of origin, 4) use local seed from natural areas 

unless another source is superior, 5) use seed from a similar 

climate if local is not available, 6) determine usefulness of indigenous and exotic species, and delimit climate zones for safe 

CHAPTER 4 • TRANSFER GUIDELINES AND ZONES • RESOURCES
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 297

to help interpret the significance of the map legend in terms of 

climate.

Link: http://gmaps.nrcan.gc.ca/climwhere/

855. Omernik JM. 1987. Ecoregions of the conterminous 

United States.

Type: Map/Government Document

Geographic Area: USA

Keywords: ecoregion, ecosystems, cartographic analysis, resource management, regional geography

Abstract: A map of ecoregions of the conterminous United 

States has been compiled to assist managers of aquatic and 

terrestrial resources in understanding the regional patterns 

of the realistically attainable quality of these resources. The 

ecoregions are based on perceived patterns of a combination 

of causal and integrative factors including land use, land surface form, potential natural vegetation, and soils. A synoptic 

approach similar to that used to define these ecoregions is 

also useful for applications of the map. Initial efforts to use 

the framework are at the state level of resource management; 

they center on aquatic ecosystems—mainly attainable ranges 

in chemical quality, biotic assemblages, and lake trophic state. 

Ecoregion maps (EPA Level III) are used by the Native Seed 

Network (NSN, http://www.nativeseednetwork.org/article_

view?id=27) to guide transfers. They recommend that in the 

absence of genetic information, this level of mapping offer a 

practical guide for seed transfer zone boundaries.

Link: http://www.epa.gov/wed/pages/ecoregions.htm

856. Oregon Department of Forestry. 2011.

Sources of native forest nursery seedlings. Portland (OR): 

Oregon Department of Forestry. 26 p.

Type: Government Document

Geographic Area: Oregon, USA

Compilers’ Keywords: seed zone maps, guidelines, manual, 

seed sources, catalog, forest management, nurseries

Introduction: Sources of Native Forest Nursery Seedlings 

is a service provided by the Oregon Department of Forestry 

to help meet the changing needs of family and non-industrial private forestland (NIPF) owners actively engaged 

in: Afforestation, Reforestation, Forest Management, Tax 

Incentives, Timber Production, Riparian Management, Forest 

Pest Management, Watershed Enhancement, Fish and Wildlife 

Habitat Improvement, Marketing of Forestry Carbon, Offsets, 

etc. Published annually, the catalog helps landowners locate 

suitable tree stock to meet their unique and individual resource 

objectives. This is just one of several valuable services that 

ODF provides to family and private non-industrial forestland 

owners. Other services include providing information and 

advice, technical assistance, and financial incentive guidance 

on: Forest Health Issues, Silvicultural Practices, and Land 

to map the similarity of potential seed collection sites across 

large regions. Considerable effort has gone into developing the required GIS database for the Great Lakes region of 

North America. Such data are becoming increasingly available in many parts of the world. Some example applications 

of Seedwhere are provided for the Great Lakes region. While 

no panacea for this complex and large problem, the Seedwhere 

tool provides a quantitative approach to visualizing the issue 

that should help resource managers.

854. McKenney DW, Mackey BG, Joyce D. 2006.

ClimWhere (formerly SeedWhere). (URL accessed 29 

November 2015) 

Type: Web Page

Geographic Area: North America

Compilers’ Keywords: climate change, seed transfer, forest 

management, seedlot selection, decision support system

About: Seedwhere is intended as a tool to support decisions 

on moving plant material across environmental gradients. 

Seedwhere is a pun on the word Software with the idea of 

“Where can I move Seed”? The model maps climate similarity across large geographic regions using the Gower similarity metric. This web-based beta-version allows users to access 

several North American wide climate models and investigate 

the similarity of current climate to projected future climates. 

The climate models used here are 300 arc seconds or approximately 10 km in resolution. While no panacea, the Seedwhere 

tool provides an intuitive approach to visualizing how far forest 

managers might risk moving seed or other plant material away 

from its maternal climate in the absence of detailed population 

genetics work. The types of questions this might be relevant 

for include: how similar are the environmental conditions at 

a seed collection site to other areas; how similar are the environmental conditions at a regeneration area to potential seed 

collection sites; how similar are the environmental conditions 

at a regeneration area to seed already in storage; how similar 

are the environmental conditions at areas where seed lots have 

been collected; where are the environmental conditions similar 

to a particular seedlot source; how similar are the environmental conditions between two particular sites; and how similar are 

the environmental conditions between two particular seedlots 

that have already been collected? Output is a color or black and 

white map that shows the similarity of climate on a 0 to 1 scale. 

Seedwhere normalizes the values for each selected climate 

variable so that the 0 to 1 similarity metric can be calculated for 

the set of selected variables. When using the climate change 

models, the program compares the current climate at the selected location to the projected climate in the selected region or 

Area of Interest—OR THE RECIPORICAL. The output also 

includes a table that summarizes the range of values of the selected climate variables over the geographic area selected. The 

table also shows what each .05 change in the Gower metric 

value means in terms of each climate variable. This is intended 

CHAPTER 4 • TRANSFER GUIDELINES AND ZONES • RESOURCES
298 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Keywords: hardwood, seed zone, adaptation, geographic information system, Tennessee

Abstract: For species that have no or limited information on 

genetic variation and adaptability to nonnative sites, there is a 

need for seed collection guidelines based on biological, climatological, and/or geographical criteria. Twenty-eight hardwood 

species are currently grown for reforestation purposes at the 

East Tennessee State Nursery. The majority of these species 

have had no genetic testing to define guidelines for seed collection location and can be distributed to sites that have a very 

different environment than that of seed origin(s). Poor survival 

and/or growth may result if seedlings are not adapted to environmental conditions at the planting location. To address this 

problem, 30 yr of Tennessee county precipitation and minimum 

temperature data were analyzed and grouped using a centroid 

hierarchical cluster analysis. The weather data and elevational 

data were entered into a Geographic Information System (GIS) 

and separately layered over Bailey’s Ecoregions to develop a 

seed zone system for Tennessee. The seed zones can be used 

as a practical guideline for collecting seeds to ensure that the 

resulting seedlings will be adapted to planting environments.

859. Randall WK, Berrang PC. 2002. Washington 

tree seed transfer zones. Olympia (WA): Washington State 

Department of Natural Resources. 84 p.

Type: Government Document

Geographic Area: Washington, USA

Compilers’ Keywords: reforestation, maladaptation, seed 

zone map, forest management, seed transfer guidelines

Introduction: Choosing the appropriate seed to reforest a 

particular site is important for many reasons: producing a longlived, healthy stand; limiting damage from climate or pests; 

promoting rapid production of commodities; and maintaining 

locally adapted gene pools. This document will provide information to land managers responsible for selecting forest tree 

seed for reforestation. The risk of moving seed from a source 

environment to a planting environment will be kept within 

acceptable levels by establishing seed zones and elevation 

bands within which seed can be transferred. These recommendations will supersede those of the Tree Seed Zone Map 

that was published for the State of Washington in 1966 (see 

next page). They apply to seed collected from natural populations of native forest trees unless otherwise noted. New tree 

seed zones or seed transfer guidelines are needed because the 

ones in current use are out of date. The old Tree Seed Zone 

Map was based only on climatic, vegetative, and topographic 

information. Now genetics information, which has accumulated over the past 30 years, needs to be incorporated into the 

recommendations. For example, the old tree seed zones were 

the same for all species, but it is now known that species differ tremendously in how far they can be moved safely. This 

guide summarizes published seed zone literature, seed transfer 

Stewardship Activities. Nurseries included in the catalog may 

have additional stock available. Please contact individual nurseries for stock information, current seedling inventories and 

ordering procedures.

Link: http://www.oregon.gov/odf/privateforests/docs/SeedlingCatalogWeb.pdf

857. Parker WH, Lesser MR. 2004.Development of focal 

point seed zones for white spruce in Northern Ontario. Thunder 

Bay, Ontario, Canada: Lakehead University. Final Report Part 

2 - Living Legacy Trust Project 04-012. 91 p.

Type: Report

Geographic Area: Ontario, Canada

Keywords: white spruce, Picea glauca, focal point seed zone, 

provenance test, adaptive variation

Abstract: White spruce (Picea glauca [Moench] Voss.) is distributed widely throughout the boreal forest of Ontario and the 

rest of Canada. Ecologically based management of this species 

requires an understanding of its patterns of adaptive variation. 

The development of focal point seed zones will assist in properly matching seed sources and planting sites. To develop focal 

point seed zones of white spruce, one hundred and thirty two 

provenances from Ontario and western Quebec were established at a greenhouse and five field trials throughout Ontario. 

Growth and phenological variables were measured over two 

growing seasons and selected variables regressed individually 

against geographic and climate variables to assess whether 

they exhibited adaptive variation. Principal components (PC) 

analysis was used to summarize the main components of 

variation. The first PC mainly represented growth potential. 

PCs 2 and 3 mainly represented phenological traits. PC factor 

scores were regressed against climate variables and the resulting equations were used to model the PC axes. Models were 

converted to spatial data and reproduced as contoured grids 

using GIS. For any given point in Ontario PC axis grids can 

be standardized and intersected, creating zones of adaptive 

similarity to that point. Focal point seed zones created for 20 

example points selected from across the province show strong 

latitudinal trends and more regional longitudinal trends. Seed 

transfer for white spruce across traditional site region boundaries may be possible in most of north-central and north-eastern 

portions of Ontario.

Link: http://flash.lakeheadu.ca/~fpszsw/LLT%20REPORT3.

pdf

858. Post LS, Schlarbaum SE, van Manen F, Cecich 

RA, Saxton AM, Schneider JF. 2003. Development of 

hardwood seed zones for Tennessee using a geographic information system. Southern Journal of Applied Forestry 27:172-175.

Type: Journal 

Geographic Area: Tennessee, USA

CHAPTER 4 • TRANSFER GUIDELINES AND ZONES • RESOURCES
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 299

Abstract: Summary of transfer guidelines for Douglas-fir in 

western Montana. Guidelines based on adaptive differentiation 

found in juvenile trees related to physiographic characteristics 

and elevation. The author details both discrete and floating 

transfer guidelines for western and eastern provinces because 

elevational clines are steeper in the west (400’) than in the east 

(800’). Small geographic and narrow elevation limits to transfer may be inappropriate, however expanding limits of transfer increases risk of maladaptation (reduced growth rate and 

increased cold damage). Regression techniques accounted for 

81% of the variance in 3-year height of 54 Douglas-fir populations from western Montana. Seed transfer guidelines are developed from patterns of adaptive variation. 

Link: http://books.google.com/

books?id=p3RAcad3EZMC&printsec=frontcover&dq=See

d+transfer+guidelines+for+Douglas-fir+in+Western+Mon

tana&hl=en&sa=X&ei=iycmUpvhF-TbigKS1ICgCQ&ved-

=0CEgQ6AEwAA#v=onepage&q=Seed%20transfer%20

guidelines%20for%20Douglas-fir%20in%20Western%20

Montana&f=false

862. Rehfeldt GE. 1987. Adaptive variation and seed transfer for ponderosa pine in Central Idaho. Ogden (UT): USDA 

Forest Service, Intermountain Research Station. RN-INT-373. 

9 p.

Type: Government Document

Geographic Area: Idaho, USA

Keywords: Pinus ponderosa, seed zones, reforestation, forest 

genetics, ecological adaptation

Abstract: When planted and compared in the same environments, populations from elevationally or geographically mild 

sites demonstrated the most growth, mainly because of the 

long duration and rapid rate of shoot elongation. Populations 

from relatively cold sites were comparatively shorter largely 

because growth ceased early. Populations from the South Fork 

of the Salmon River, however, combined a high rate of shoot 

elongation with a short duration of growth and thereby were 

capable of high productivity while maintaining adaptability for 

severe sites. In artificial reforestation, seed transfer from most 

populations should be limited to ± 650 feet of the elevation at 

the source. Test should be established to determine if South 

Fork populations can be moved beyond the recommended limits in an attempt to increase productivity on harsh, cold sites. 

Link: http://books.google.com/books?id=lbhLB80od2cC&p

rintsec=frontcover&source=gbs_ge_summary_r&cad=0#v=

onepage&q&f=false

863. Riley LE, Dumroese RK, Landis TD. 2002.

National proceedings: forest and conservation nursery associations—2002. Ogden (UT): USDA Forest Service, Rocky 

Mountain Research Station. RMRS-P-28. 187 p. 

Type: Government Document

rules, genetics, and geographic variation for tree species used 

in reforestation, wildlife, and riparian planting. Specific guidelines are given for each species. These guidelines are meant to 

improve silvicultural prescriptions, not to replace them. Not 

all sites within a seed zone will be appropriate for a particular tree species. You must rely on your knowledge of species 

characteristics to determine which species is most appropriate 

for the site you plan to reforest. To determine the best source 

of seed for the area you want to plant, locate the page reference 

for that species in the table of contents and read the specific 

recommendation. Then refer to the species map for seed zones 

and elevations bands. Elevation bands are not mapped, but are 

considered in the seed transfer guidelines. Also, seed zones are 

only delineated for areas where the species naturally occurs. 

Each of these guidelines is for a particular forest tree species 

and should not be used for other plants. However, the 1966 

Tree Seed Zones encompass areas where environmental variation is fairly uniform and could serve as guidelines for other 

species where no seed zones have been established.

Link: http://www.dnr.wa.gov/ResearchScience/Topics/ForestResearch/Pages/lm_tree_seed_zones.aspx

860. Rehfeldt GE. 1980. Seed transfer guidelines for 

Douglas-fir in North Idaho. Ogden (UT): USDA Forest Service, 

Intermountain Forest and Range Experiment Station. RNINT-300. 4 p.

Type: Government Document

Geographic Area: Idaho, USA

Compilers’ Keywords: seed transfer guidelines, Pseudotsuga 

menziesii, maladaptation, genetic adaptation, reforestation, 

delineations

Compilers’ Summary: Seed transfer guidelines for Douglasfir reforestation in Northern Idaho. Includes delineations based 

on elevation and latitude using current research (1980). Author 

mentions that there will be some maladaptation in planting 

even when adhering to the guidelines, but as more information 

becomes available through research they can be fine-tuned. 

Because patterns of variation in Douglas-fir are so high, these 

guidelines should be followed in plantings. 

Link: http://books.google.com/books?hl=en&lr=&id=7yis1

cDGdbUC&oi=fnd&dq=Seed+transfer+guidelines+for+Do

uglas-fir+in+North+Idaho&ots=Hmq45XH7XG&sig=7PdS

wGR8grJyKXMHgQy6pD67yFI

861. Rehfeldt GE. 1983. Seed transfer guidelines for 

Douglas-fir in Western Montana. Ogden (UT): USDA Forest 

Service, Intermountain Forest and Range Experiment Station. 

RN-INT-329. 5 p.

Type: Government Document

Geographic Area: Montana, USA

Keywords: seed zones, seed transfer, adaptive variation

CHAPTER 4 • TRANSFER GUIDELINES AND ZONES • RESOURCES
300 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

Compilers’ Keywords: restoration, genetic x environment interaction, seed transfer guidelines

Abstract: Atriplex canescens (Pursh.) Nutt. is the most 

widespread species of perennial Atriplex in North America. 

Throughout its distributional range, A. canescens shows considerable between-population variation. Some of this variation 

may be due to phenotypic plasticity but most of it appears to 

be genetic. Mutations, polyploidy, introgressive hybridization, 

and segregation from interspecific hybrids all appear to have 

contributed to its extensive heritable variation. Polyploidy is 

unusually common with numerous chromosome races (2x, 4x, 

6x, 8x, 10x, 12x, 14x, 20x). Fourwing saltbush is widely used 

for reclamation plantings. Proper identification is important to 

the utilization of fourwing saltbush in such plantings. While 

many of the races have been formally named as varieties, others have not. Even though differentiated by ploidy, chemical 

constituents, geographic distribution, and statistical distribution of morphological characters, races may lack sufficient 

diagnostic characters to allow facile identification, at least 

in the herbarium. Rather than combining named races under 

those that do have a taxonomic name, it seems better at present not to use the formal intraspecific categories in treating the 

fourwing saltbushes, but to consider them all as races. Seed 

transfer should be within the geographical distribution limits 

of each race. The most common race, by far, is Occidentalis. 

We recommend four overlapping seed transfer zones for race 

Occidentalis in the United States: (1) Northern Intermountain, 

(2) Western Great Plains, (3) Colorado Plateau/ Great Basin/

Columbia Basin, and (4) Southwestern. Source seed populations from near the planting sites generally do well; and 

populations generally perform better when moved south and/

or to lower elevations than when moved north and/or up in 

elevation.

Link: http://www.treesearch.fs.fed.us/pubs/6273

866. Schmidtling RC. 2001. Southern pine seed sources. Asheville (NC): USDA Forest Service, Southern Research 

Station. General Technical Report GTR-SRS-44. 35 p.

Type: Government Document

Geographic Area: Southeastern USA

Keywords: fusiform rust, geographic variation, loblolly pine, 

longleaf pine, provenance tests, sand pine, seed movement, seed 

sources, shortleaf pine, slash pine, Virginia pine

Abstract: The selection of an appropriate seed source is critical for successful southern pine plantations. Guidelines for selection of seed sources are presented for loblolly (Pinus taeda

L.), slash (P. elliottii Engelm.), longleaf (P. palustris Mill.), 

Virginia (P. virginiana Mill.), shortleaf (P. echinata Mill.), and 

sand [P. clausa (Chapm. ex Engelm.) Vasey ex Sarg.] pines. 

Seed movement guidelines in this handbook are based on climatic similarities between the seed source origin and the planting site. Because yearly average minimum temperature is the 

Geographic Area: USA

Keywords: bareroot nursery, container nursery, nursery practices, fertilization, pesticides, seeds, reforestation, restoration, 

plant propagation, native plants, tree physiology

Abstract: This proceedings is a compilation of 33 papers that 

were presented at the regional meetings of the forest and conservation nursery associations in the United States in 2002. The 

joint meeting of the Southern Forest Nursery Association and 

the Northeastern Forest and Conservation Nursery Association 

was held at the DoubleTree Hotel and Conference Center in 

Gainesville, Florida on July 15 to 18. The meeting was hosted 

by the Florida Division of Forestry, Andrews Nursery. In addition to technical sessions, tours included Andrews Nursery 

and Stansel Farm & Nursery. The combined meeting of the 

Western Forest and Conservation Nursery Association and 

the Forest Nursery Association of British Columbia occurred 

August 5 to 8 at the WestCoast Hotel in Olympia, Washington. 

The meeting was hosted by the Washington Department of 

Natural Resources, Webster Forest Nursery. Morning technical 

sessions were followed by field trips to Webster Nursery and 

the Weyerhaeuser Bonsai Garden. Subject matter for both sessions included seed transfer, collection, and processing; pest 

problems and pesticide use; nursery culturing; transplanting; 

harvesting, storage, and outplanting.

Link: http://www.fs.fed.us/rm/pubs/rmrs_p028.pdf

864. Saari C, Glisson W, Borzym N, Sotern 

B. 2011. Chicago region restoration seed source policies.

Chicago (IL): Northwestern University. 30: 3. 59 p.

Type: Report

Geographic Area: Illinois, USA

Compilers’ Keywords: seed source selection, native plants, 

climate change, provenance, decision-making, restoration

Abstract: This report seeks to summarize seed source policies 

and practices as understood and implemented by the conservation agencies and organizations of the Chicago region. We 

also consider current scientific research on seed provenance 

and discuss some approaches that take into account both the 

ongoing research as well as considerations suggested by global 

climate change. Our goal is to provide information and analysis that may be helpful to decision-making for seed provenance 

in ecosystem restoration.

Link: http://www.habitatproject.org/webdocs/misc/ChicagoRegionSeedSourcePolicies.pdf

865. Sanderson SC, McArthur ED. 2004. Fourwing 

saltbush (Atriplex canescens) seed transfer zones. Fort Collins 

(CO): USDA Forest Service, Rocky Mountain Research Station. 

General Technical Report RMRS-GTR-125. 15 p.

Type: Government Document

Geographic Area: Western USA

CHAPTER 4 • TRANSFER GUIDELINES AND ZONES • RESOURCES
USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015 301

Type: Government Document

Geographic Area: Oregon, USA

Keywords: Pinus ponderosa, seedlings, provenance, geographic genetic variation, ecological genetics, genecology, genetic 

diversity

Abstract: Adaptive genetic variation in seed and seedling 

traits for ponderosa pine from the east slopes of the Cascade 

Range in Oregon was analyzed by using 307 families from 

227 locations. Factor scores from three principal components 

based on seed and seedling traits were related by multiple regression to latitude, distance from the Cascade crest, elevation, 

slope, and aspect of the seed source and by classification analysis to seed zone and 300-meter elevation band within zone. A 

provisional transfer risk equation and tentative new seed zones 

were delineated to guide seed transfer in artificial regeneration.

Link: http://www.treesearch.fs.fed.us/pubs/4989

870. Sorensen FC, Weber JC. 1994. Genetic variation 

and seed transfer guidelines for ponderosa pine in the Ochoco 

and Malheur National Forests of Central Oregon. Portland (OR): 

USDA Forest Service, Pacific Northwest Region. Research 

Paper PNW-RP-468. 36 p.

Type: Government Document

Geographic Area: Oregon, USA

Keywords: Pinus ponderosa, seedlings, breeding zones, provenance, genetic variation, adaptive variation, genetic diversity

Abstract: Adaptive genetic variation in seed and seedling 

traits was evaluated for 280 families from 220 locations. 

Factor scores from three principal components were related 

by multiple regression to latitude, longitude, elevation, slope, 

and aspect of the seed source, and by classification analysis to 

seed zone and elevation band in seed zone. Location variance 

was significant but not large. Multiple regression equations explained less than 50 percent of location variance. Slope aspect 

variables were important.

Link: http://www.treesearch.fs.fed.us/pubs/20581

871. St. Clair JB, Johnson RC, Shaw NL. 2011.

Genecology and seed transfer guidelines for bluebunch wheatgrass (Pseudoroegneria spicata). In: 2011 Great Basin Native 

Plant Selection and Increase Project Annual Meeting. 27 p.

Type: Presentation

Geographic Area: Western USA

Compilers’ Keywords: genetic variation, cultivar, native 

bunchgrass, provenance test

Compilers’ Summary: Presentation from the 2011 Great 

Basin Native Plant Selection and Increase Project Annual 

Meeting on the genecology and seed transfer guidelines for 

bluebunch wheatgrass to identify adapted and diverse populations for restoration. Using collections from 127 populations 

most important climatic variable related to growth and survival, it has been used to define the rules of seed movement. This 

variable, which defines plant hardiness zones, has been used 

for many years by horticulturists to guide the transfer of plant 

materials. East-west movement to areas of similar climate is 

permissible, with the exception of loblolly pine.

Link: http://www.treesearch.fs.fed.us/pubs/2797

867. Schubert GH, Pitcher JA. 1973. A provisional 

tree seed-zone and cone-crop rating system for Arizona and 

New Mexico. Fort Collins (CO): USDA Forest Service, Rocky 

Mountain Research Station. RM-105. 15 p.

Type: Government Document

Geographic Area: Southwestern USA

Keywords: cone collecting, forest seed collecting, forest seed 

production

Abstract: The forested areas of Arizona and New Mexico 

were divided into 10 physiographic-climatic regions. These 

regions were then subdivided into five to nine seed collection 

zones about 50 miles wide. Provenance tests will be conducted 

to determine variation and need for adjustments. Seed used for 

reforestation should be limited to that collected within the local 

zone. A 10-unit classification system for rating cone crops is 

included.

Link: http://search.library.wisc.edu/catalog/ocm05165931

868. Sorensen FC. 1992.Genetic variation and seed transfer 

guidelines for lodgepole pine in central Oregon. Corvallis (OR): 

USDA Forest Service, Pacific Northwest Research Station. 

PNW-RP-453. 36 p.

Type: Government Document

Geographic Area: Oregon, USA

Keywords: Pinus contorta var. murrayana, lodgepole pine, 

geographic variation, seed transfer, breeding zones

Abstract: Cones were collected from 272 trees at 189 locations uniformly distributed over the east slopes of the Oregon 

Cascade Range and Warner Mountains. Variation in seed and 

seedling traits was related to (1) seed source latitude, distance 

from the Cascade crest, elevation, slope, and aspect in multiple 

regression analyses; and (2) seed zone and elevation band in 

classification analyses. Provisional seed transfer guidelines 

are presented. These include a regression equation for guiding 

seed transfer and estimating transfer risk, and a new outline of 

fixed seed zones.

Link: http://www.treesearch.fs.fed.us/pubs/20568

869. Sorensen FC. 1994.Genetic variation and seed transfer 

guidelines for ponderosa pine in Central Oregon. Corvallis (OR): 

USDA Forest Service, Pacific Northwest Research Station. 

PNW-RP-472. 30 p.

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302 USDA Forest Service Gen. Tech. Rep. RMRS-GTR-347. 2015

About: It’s easy to support a motherhood statement like “genetic diversity should be conserved.” After more than a decade of attention to this topic, most of us realize that genetic 

variation provides the raw material for future selection in tree 

improvement programs for new traits (e.g., resistance to new 

insects or diseases, or changes in fiber quality to meet new 

industrial demands). We also realize that natural populations 

require genetic diversity to adapt to new environmental conditions, to allow evolution to proceed. But how do we go about 

conserving genetic diversity, and how can we rigorously assess 

whether we are meeting this goal? In 2000, the Forest Genetics 

Council of British Columbia (FGC) realized that, while gene 

conservation continued to be a high priority, this objective 

was not being met in a strategic and rigorous manner. As a 

result, the Centre for Forest Conservation Genetics (CFCG) 

was established in the Department of Forest Sciences at the 

University of British Columbia (UBC). Funding is provided 

from the Forest Investment Account (FIA). The CFCG has a 

mandate from the Forest Genetics Council to (1) study population genetic structure of forest trees using existing or new data; 

(2) assess the current degree of gene conservation both in situ 

in existing reserves and ex situ collections, and the need for 

additional protection; and (3) evaluate the current degree of 

maintenance of genetic diversity in breeding and deployment 

populations of improved varieties to meet current and future 

environmental challenges. This website is a portal for forest 

genetics and climate change research conducted in British 

Columbia, Canada.

Link: http://www.genetics.forestry.ubc.ca/cfcg/

874. University of British Columbia. 2010.

AdapTree. Vancouver, British Columbia, Canada: 

University of British Columbia, Department of Forest 

Sciences, Centre for Forest Conservation Genetics. (URL 

accessed 29 November 2015)

Type: Web Page

Geographic Area: Western Canada

Compilers’ Keywords: climate change, seed transfer, forest 

management, genomics

About: The primary objective of this project is to improve 

provincial seed transfer policy and operational forest management response to climate change by: comparing the adaptive 

portfolio of operational seedlots from tree breeding programs 

and seed orchards to the climatic distribution and landscape 

genomics of natural populations; developing strategies for operational seed transfer that will reduce the risk of loss of forest 

productivity and health due to maladaptation in planted forests; evaluating ecological, economic, social, and legal implications of these results for forest-dependent communities and 

ecosystems.

Link: http://adaptree.sites.olt.ubc.ca/

growing in Washington, Oregon, California, Nevada, Utah, 

Idaho and Montana and common gardens established in 3 test 

sites, the authors found large differences among test sites and 

between years in size and phenology, high levels of population 

variation in many traits, population means strongly correlated 

among test sites and years, size traits intercorrelated, phenology traits intercorrelated, other traits not correlated with each 

other, and strong correlations with climate. There is strong evidence for adaptively significant genetic variation in bluebunch 

wheatgrass, which supports the need to delineate seed zones 

along temperature and moisture (aridity) gradients. 

Link: http://www.fs.fed.us/rm/boise//research/shrub/projects/PowerPoint_Presentations/2011/stclair.pdf

872. St. Clair JB, Howe GT, Wright JW, Beloin R. 

2011. Center for forest provenance data. Corvallis (OR): 

Oregon State University and USDA Forest Service. (URL accessed 29 November 2015)

Type: Web Page

Geographic Area: North America

Compilers’ Keywords: provenance test, forest, genetic research, online database, climate change

About: The Center for Forest Provenance Data is a place for 

researchers to archive provenance data and genecology studies of forest trees and make available for collaboration with 

other researchers. Studies consider genetic variation among 

forest trees from different source locations by growing them in 

replicated tests in a common environment such that observed 

differences are primarily due to genotype and not the environment. Consistent differences among sources that are associated 

with environmental gradients are indicative of adaptively significant variation. These studies are important for understanding genetic variation across the landscape and for managing 

genetic resources for reforestation, restoration, gene conservation, and responding to climate change. The Center for Forest 

Provenance Data has sections for submitting and retrieving 

data from the database. There is also a search tool for determining studies that are currently in the database. Currently, the 

database includes provenance information for ponderosa pine, 

Douglas-fir, whitebark pine, sugar pine, and white fir. 

Link: http://cenforgen.forestry.oregonstate.edu/index.php

873. University of British Columbia. 2000. Centre 

for forest conservation genetics. Vancouver, British Columbia, 

Canada: University of British Columbia, Department of Forest 

Sciences. (URL accessed 29 November 2015)

Type: Web Page

Geographic Area: Western Canada

Compilers’ Keywords: adaptation, forest management, tree 

populations, diversity, environmental change

CHAPTER 4 • TRANSFER GUIDELINES AND ZONES • RESOURCES
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Compilers’ Keywords: genetics, tree improvement

Introduction: The present boom in forest genetics makes it 

difficult for even the specialist to keep abreast of all the latest 

developments in this field. Therefore, in the fall of 1954 the 

committee on Forest Tree Improvement, Society of American 

Foresters, asked the author to prepare a bibliography of publications issued in 1954. As far as possible, all references that 

have a bearing on the genetics of forest trees are included. 

Also, some borderline references (for example, those on vegetative propagation) are included because they are primarily of 

interest to tree breeders. This bibliography was prepared from 

standard forestry, botanical, and abstracting journals; experiment station publications; and correspondence with tree breeders in this country and abroad. The bibliography is arranged 

alphabetically by authors. A subject-matter index can be found 

at the end. 

Link: http://www.treesearch.fs.fed.us/pubs/13626

875. USDA Forest Service. 2008. FSM 2000—National 

Forest Resource Management - Vegetation Ecology. Washington 

(DC): USDA Forest Service. FSM 2000 Chapter 2070. 12 p.

Type: Government Document

Geographic Area: USA

Compilers’ Keywords: native plant materials, restoration, 

vegetation, ecology, diversity

Compilers’ Synopsis: USFS manual for vegetation ecology 

and include authority and law pertaining to management and use 

of native and non-native plant materials on NFS lands and other 

lands under Forest Service administration. 

Link: http://www.fs.fed.us/cgi-bin/Directives/get_dirs/

fsm?2000!

876. Western Wildland Environmental Threat 

Assessment Center. 2012. Seed zone mapper. Prineville 

(OR): USDA Forest Service. (URL accessed 29 November 

2015)

Type: Web Page

Geographic Area: North America

Compilers’ Keywords: seed zone delineation, tool, GIS, climate change, restoration, native plant transfer

About: Seed zone mapper is a mapping and planning tool 

for plant material development, gene conservation, and native 

plant restoration. A seed zone is an area within which plant materials can be transferred with little risk of being poorly adapted to their new location. This site allows end-users to view and 

acquire data on seed zones for use in plant material development, gene conservation and native plant restoration activities. 

Users can also evaluate seed zones in relation to other map 

services and wildland threats published by WWETAC such as 

climate change projections or wildfire risk. Client applications 

range in functionality from a simple geobrowser (requires only 

a web browser) to ArcGIS ArcMap, a full-feature GIS software 

platform that allows the user to integrate their own data and 

create map layouts. These seed zone mapping applications are 

part of a family of Wildland Threat Mapping (WTM) applications developed by WWETAC to portray the spatial interactions of wildland threats and high value resources that occur 

in wildlands. Visit WWETAC’s WTM page for a collection of 

these mapping applications.

Link: http://www.fs.fed.us/wwetac/threat_map/

SeedZones_Intro.html

877. Wright JW. 1957. A bibliography on forest genetics and forest tree improvement 1955. Upper 

Darby (PA): USDA Forest Service, Northeastern Forest 

Experiment Station. Station Paper NE-90. 60 p.

Type: Government Document

Geographic Area: North America

CHAPTER 4 • TRANSFER GUIDELINES AND ZONES • RESOURCES
United States Department of Agriculture

Forest Service

Rocky Mountain Research Station

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gender identity (including gender expression), sexual orientation, disability, age, marital status, family/parental 

status, income derived from a public assistance program, political beliefs, or reprisal or retaliation for prior civil 

rights activity, in any program or activity conducted or funded by USDA (not all bases apply to all programs). 

Remedies and complaint filing deadlines vary by program or incident.

Persons with disabilities who require alternative means of communication for program information (e.g., 

Braille, large print, audiotape, American Sign Language, etc.) should contact the responsible Agency or USDA’s 

TARGET Center at (202) 720-2600 (voice and TTY) or contact USDA through the Federal Relay Service at 

(800) 877-8339. Additionally, program information may be made available in languages other than English.

To file a program discrimination complaint, complete the USDA Program Discrimination Complaint Form, 

AD-3027, found online at http://www.ascr.usda.gov/complaint_filing_cust.html and at any USDA office or 

write a letter addressed to USDA and provide in the letter all of the information requested in the form. To 

request a copy of the complaint form, call (866) 632-9992. Submit your completed form or letter to USDA by: 

(1) mail: U.S. Department of Agriculture, Office of the Assistant Secretary for Civil Rights, 1400 Independence 

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