Macroecology, Paleoecology, and

Ecological Integrity of Terrestrial

Species and Communities of the

Interior Columbia Basin and

Northern Portions of the Klamath

and Great Basins

United States

Department of

Agriculture

Forest Service

Pacific Northwest

Research Station

United States

Department of the

Interior

Bureau of Land

Management

General Technical

Report

PNW-GTR-410

October 1998 B.G. Marcot, L.K. Croft, J.F. Lehmkuhl, R.H. Naney, C.G. Niwa,

W.R. Owen, and R.E. Sandquist
Authors

B.G. MARCOT is a wildlife ecologist, Pacific Northwest Research Station, Forestry Sciences

Laboratory, P.O. Box 3890, Portland, OR 97208-3890; L.K. CROFT is a plant ecologist, Ochoco

National Forest, P.O. Box 490, Prineville, OR 97754; J.F. LEHMKUHL is a research wildlife

biologist, Pacific Northwest Research Station, Forestry Sciences Laboratory, 1133 N. Western

Avenue, Wenatchee, WA 98801; R.H. NANEY is a wildlife biologist, Okanogan National Forest,

1240 S. Second, Okanogan, WA 98840; C.G. NIWA is a research entomologist, Pacific Northwest

Research Station, Forestry Sciences Laboratory, 3200 S.W. Jefferson Way, Corvallis, OR 97331; W.R.

OWEN is a forest botanist, Boise National Forest, 1750 Front Street, Boise, ID 83702; and R.E.

SANDQUIST is a regional entomologist, Pacific Northwest Region, P.O. Box 3623, Portland, OR

97208-3623.
Macroecology, Paleoecology, and Ecological

Integrity of Terrestrial Species and

Communities of the Interior Columbia River

Basin and Northern Portions of the Klamath

and Great Basins

B.G. Marcot, L.K. Croft, J.F. Lehmkuhl, R.H. Naney,

C.G. Niwa, W.R. Owen, and R.E. Sandquist

Interior Columbia Basin Ecosystem Management Project:

Scientific Assessment

Thomas M. Quigley, Editor

U.S. Department of Agriculture

Forest Service

Pacific Northwest Research Station

Portland, Oregon

General Technical Report PNW-GTR-410

October 1998
Abstract

Marcot, B.G.; Croft, L.K.; Lehmkuhl, J.F.; Naney, R.H.; Niwa, C.G.; Owen, W.R.; Sandquist,

R.E. 1998. Macroecology, paleoecology, and ecological integrity of terrestrial species and communities of the interior Columbia River basin and northern portions of the Klamath and Great

Basins. Gen. Tech. Rep. PNW-GTR-410. Portland, OR: U.S. Department of Agriculture, Forest

Service, Pacific Northwest Research Station. 131 p. (Quigley, Thomas M., tech. ed. Interior

Columbia River Basin Ecosystem Management Project: scientific assessment).

This report presents information on biogeography and broad-scale ecology (macroecology) of selected

fungi, lichens, bryophytes, vascular plants, invertebrates, and vertebrates of the interior Columbia River

basin and adjacent areas. Rare plants include many endemics associated with local conditions. Potential

plant and invertebrate bioindicators are identified. Species ecological functions differ among communities and variously affect ecosystem diversity and productivity. Species of alpine and subalpine communities are identified that may be at risk from climate change. Maps of terrestrial ecological integrity

are presented.

Keywords: Macroecology, paleoecology, ecological integrity, terrestrial communities, ecosystems,

wildlife, fungi, lichens, bryophytes, vascular plants, invertebrates, arthropods, mollusks, amphibians,

reptiles, birds, mammals, endemism, interior Columbia River basin, Klamath Basin, Great Basin.
Preface

The Interior Columbia Basin Ecosystem Management Project was initiated by the Forest Service and

the Bureau of Land Management to respond to several critical issues including, but not limited to, forest

and rangeland health, anadromous fish concerns, terrestrial species viability concerns, and the recent

decline in traditional commodity flows. The charter given to the project was to develop a scientifically

sound, ecosystem-based strategy for managing the lands of the interior Columbia River basin administered by the Forest Service and the Bureau of Land Management. The Science Integration Team was

organized to develop a framework for ecosystem management, and assessment of the socioeconomic

and biophysical systems in the basin, and an evaluation of alternative management strategies. This

paper is one in a series of papers developed as background material for the framework, assessment, or

evaluation of alternatives. It provides more detail than was possible to disclose directly in the primary

documents.

The Science Integration Team, although organized functionally, worked hard at integrating the approaches, analyses, and conclusions. It is the collective effort of team members that provides depth and

understanding to the work of the project. The Science Integration Team leadership included deputy

team leaders Russel Graham and Sylvia Arbelbide; landscape ecology—Wendel Hann, Paul Hessburg,

and Mark Jensen; aquatic—Jim Sedell, Kris Lee, Danny Lee, Jack Williams, Lynn Decker; economic—

Richard Haynes, Amy Horne, and Nick Reyna; social science—Jim Burchfield, Steve McCool, and

Jon Bumstead; terrestrial—Bruce Marcot, Kurt Nelson, John Lehmkuhl, Richard Holthausen, and

Randy Hickenbottom; spatial analysis—Becky Gravenmier, John Steffenson, and Andy Wilson.

Thomas M. Quigley

Editor

United States

Department of

Agriculture

Forest Service

United States

Department of

the Interior

Bureau of Land

Management

Interior Columbia

Basin Ecosystem

Management Project
Executive Summary

Plant Biodiversity

Biodiversity of the interior Columbia River basin assessment area (hereafter referred to as the basin

assessment area) is better known than is global biodiversity, but much systematic and inventory work

remains to be done on soil micro-organisms, fungi, and invertebrates. Rare plants and lichen groups

occur throughout the basin assessment area, and rare plants were particularly diverse in the Basin and

Range, Columbia Plateau, Blue Mountains, northern Idaho, and the southern regions of the east side of

the Cascade Range. Most rare plant life forms were hemicryptophytes and cryptophytes. Diversity of

rare plants was not particularly oriented to specific topographic conditions, as many rare plants occur

in various mostly azonal (atypical) conditions. Most rare plants are declining with fewer remaining

stable; by definition, none is significantly increasing. Many paleoendemic (ancient endemic) and

neoendemic (recently endemic) plants occur in the basin assessment area. Lichens disperse mostly

by gravity; rare fungi by wind or vertebrates; and rare vascular plants by gravity, wind, water, and

vertebrates. Most of the rare plants are pollinated probably by various invertebrates. Several lichens,

bryophytes, plants, and invertebrates can serve as bioindicators of environmental conditions and

ecosystem health.

Animal Biodiversity

Invertebrates of the basin assessment area disperse by many means. One way involving coevolution

between invertebrates and vertebrates is phoresis (dispersal by hitching a ride on vertebrate organisms).

Disease and parasites are poorly studied in the basin assessment area and need further work to determine distribution, frequency of occurrence, and effects on plants and animals of interest to management. Endemic vertebrates include some amphibians, birds, and mammals. The ranges of many vertebrate species extend beyond the basin assessment area, including semiendemics (species occurring

only within the inland West or the basin assessment area for only a portion of the year, as with some

migrants). Thus, to provide for their rangewide viability, conditions beyond the basin assessment area

need to be known and addressed. Numbers of breeding birds by state match other published predictions.

Most bird species were ranked as common, with fewer uncommon, rare, abundant, or irregular. More

abundant and irregularly occurring bird species occur toward midcontinent than elsewhere in the basin

assessment area. Most birds are resident or summer breeders, with fewer being migrants or wintering

only in the basin assessment area.

Ecological Functions of Species Among Communities

We present species function profiles that relate key ecological functions (KEFs)—those major roles

that each species plays in their environment—of vertebrates to occurrence by vegetation community.

Key ecological functions of species can affect ecosystem productivity, diversity, and sustainability;

and function profiles of species help summarize the diversity of species functions in ecological communities of the basin assessment area. Function profiles of species display the degree of functional redundancy (number of species sharing the same KEF) within each community. Some functions, particularly cavity excavation in snags and primary burrow excavation in soil, are highly variable among communities in terms of number of species. Late seral forests provide the greatest redundancy in carrion

feeding, general nutrient cycling, and primary cavity excavation functions of vertebrates. We discuss

functions and functional redundancy of other communities as well. We pose several hypotheses relating

species functional redundancy to ecosystem resiliency and changes in species community structure, as

one facet of ecosystem management. The greatest vertebrate functional diversity is found in early seral

montane forest, followed by upland woodlands and riparian woodlands, and upland shrublands. No one

community, however, contains all ecological functions of species.
Effects of Climate Change

Near-future changes in regional climates from human activities may be complex, involving increasing

interseasonal and interannual variations in precipitation and temperature. Species preadapted to such

variations likely will persist during climate changes. The fate of vertebrates within alpine tundra and

subalpine forests may differ individualistically according to latitudinal shifts of these habitats and the

range of plasticity of each species. Studies of climate change suggest that patterns of climate, vegetation, and plant and animal species occurrence in the basin assessment area during historic (since 1800)

times do not match those over the past four centuries or prehistorically during the Quaternary or Tertiary periods. This has important ramifications for reinterpreting the range of natural historic conditions

(variations), which do not represent conditions under which species evolved.

Prehistoric Conditions

The assemblage of mammalian taxa of the basin assessment area has changed greatly since Tertiary

and Quaternary periods, in accord with changes in climate and vegetation, although overall diversity

of species among families and orders has remained more or less constant. Many species, and some

families and orders, have declined or become extinct, whereas others have radiated and grown. We

anticipate future losses and gains in species, families, and orders of plants and animals over evolutionary time, although prehistoric trends in extinction and speciation (evolution of new species) do

not represent short-term ecological conditions.

Ecological Integrity of Terrestrial Communities

We mapped various aspects of terrestrial ecological integrity, including components of species viability

(as threatened, endangered and candidate species, locally endemic species, distribution of rare plants,

and key habitat corridors); long-term evolutionary potential of species (as disjunct populations and biodiversity and endemism hot spots); and multiple ecological domains (as peripheral species and unique

species assemblages, and the full set of large vertebrate carnivores). Results are shown by watershed.
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Introduction

History and Impetus

The Science Integration Team of the Interior

Columbia Basin Ecosystem Management Project

(ICBEMP) conducted ecological assessments

during 1993-96 as a joint venture among USDA

Forest Service (FS), USDI Bureau of Land Management (BLM), and many cooperators. The Science Integration Team developed the first comprehensive database and assessment of terrestrial

plant and animal ecology of the interior Columbia

River basin and northern portions of the Klamath

and Great Basins in the United States (collectively, and hereafter, referred to as the basin

assessment area) (Marcot and others 1997).

This document expands on findings related to

macroecology and ecological integrity of terrestrial species and communities of the basin

assessment area.

Our ultimate aim is to contribute to a scientific

foundation for ecosystem management of terrestrial ecosystems in the basin assessment area.

According to the Ecological Society of America

(1995), a scientific basis for ecosystem management should provide adequate information on biological diversity and on the function and dynamics of ecosystems, and acknowledge the “openness and interconnectedness” of ecosystems.

Other facets of ecosystem management should

entail study of past and current natural ranges of

conditions (Morgan and others 1994) and the

ecological roles of species interactions (Willson

1996). This report provides information on these

topics in terrestrial ecosystems of the basin

assessment area.

Definitions

In this report, macroecology refers to habitat and

environmental characteristics of individual terrestrial species or species groups at the broad

scale (sensu Brown 1995). Terrestrial species

refers to species whose life histories are mostly

confined to nonaquatic environments, although

some largely aquatic vertebrate species such as

amphibians and otters (but not fish) are included.

Hydrologic unit codes (HUCs), as used in the

ICBEMP ecology assessments, are standard delineations of watershed and subwatershed land areas

based on river drainage patterns (Jensen and

Bourgeron 1994). We used two levels of HUCs:

4th- and 6th-level hydrologic units, corresponding

to subbasins and subwatersheds, respectively.

Broad scale is defined here as such basin assessment area-wide depictions that use characterizations of 1-km2 cells with summarization by

4th-code hydrologic units. Mid scale is defined

as characterizations at 4-ha resolution with summarization by 6th-code hydrologic units.

CHAPTER 1

1
Study Area

Area and Boundaries

The basin assessment area encompasses

58 361 400 ha. It is confined on the north by the

United States-Canada border; on the east by the

Continental Divide; on the west by the crest of

the Cascade Range; on the south by the OregonCalifornia and Oregon-Nevada borders; and

elsewhere by the extent of the Columbia River

drainage (fig. 1). One major portion of the

Columbia River basin in the United States is

the tributary Snake River basin, which includes

central and southern Idaho and western Montana.

It extends toward the Continental Divide into the

Greater Yellowstone Ecosystem and southern

Yellowstone National Park. The entire Columbia

River basin encompasses about 67 466 000 ha

and extends north into British Columbia, Canada,

(15 percent of the entire area of the Columbia

River basin proper) and west past the crest of the

Cascade Range to the Pacific Ocean, outside the

boundaries of the basin assessment area (fig. 1).

The basin assessment area was defined by the directors of the ICBEMP according to both agency

planning needs and hydrologic (watershed) criteria. The existing Northwest Forest Plan and its

preceding ecological assessment (Forest Ecosystem Management Assessment Team [FEMAT]

1993) had already addressed the area within the

range of the northern spotted owl, including the

eastern slope of the Cascade Range. The ICBEMP

was directed to reevaluate the FEMAT assessments along the east side of the Cascade Range,

given the different composition of forests found

there as compared to forests west of the crest of

the Cascade Range.

The basin assessment area includes portions of

seven Western states, including major portions of

Washington, Oregon, and Idaho, as well as western Montana and corners of Wyoming, Utah, and

Nevada. Although outside the Columbia River

basin per se, the northern portions of the Klamath

Basin (1.5 million ha of the basin assessment

area) and Great Basin (4.2 million ha of the basin

assessment area) in southern Oregon were included in the basin assessment area. The purpose

was to complete broad-scale land planning and

assessments on BLM and FS lands within the

regional boundaries of these agencies in Oregon.

The portions of the Klamath Basin and northern

Great Basin are therefore also included in the

assessments of this report, although they do not

form a discrete ecological unit.

Figure 1—Boundaries of the Columbia River basin in

North America (shaded area), and the basin assessment area (heavy solid line) and Western States (light

solid lines) in the United States.

CHAPTER 2

2
By Land Ownership

Bureau of Land Management and FS lands constitute 53 percent of the land within the basin

assessment area, with the remainder in private

(38 percent), State and other Federal (4 percent),

tribal (4 percent), and national park and other

non-Federal wilderness (1 percent) lands (fig. 2).

The basin assessment area includes portions of

major ecosystems of interest to Federal broadscale planning concerns, including the Greater

Yellowstone Ecosystem, the mountain ranges of

the North Cascades/Bitterroots, the Selkirks, the

Cabinet-Yaks, the High Cascades, the Klamath

Basin, the Great Basin, and others.

By Type of Management

Most of the basin assessment area is roaded and

actively managed for natural resource use (fig. 3).

Of all ownerships within the basin assessment

area, 81 percent of the area is roaded and actively

managed (43 percent BLM-FS plus 38 percent

other lands), 10 percent is unroaded or not actively managed (9 percent BLM-FS plus 1 percent National Park Service and other wilderness

lands), and 10 percent is tribal, state, and other

public lands of varying use, mostly roaded and

actively managed. Of all BLM and FS lands

within the basin assessment area, 83 percent is

roaded and actively managed for natural resource

use and extraction, and 17 percent is characterized as unroaded.

Ecological Reporting Units

The Science Integration Team mapped 13 ecological reporting units (ERUs) in the basin assessment area. Ecological reporting units are geographically defined areas identified by the Science Integration Team by use of multiple landscape and watershed criteria and were based

on subsection delineations of 6th-code HUCs

(Jensen and others, in prep.). Ecological reporting

units represent large geographic areas with common landforms and drainage patterns, such as the

Upper Snake headwaters, northern glaciated

mountains, and the east side of the Cascade

Range in Oregon.

Hydrologic Delineations and

Biophysical Characterizations

The basin assessment area was partitioned into

164 subbasins or 4th-code HUCs, and 7,733 subwatersheds or 6th-code HUCs. The subbasins

averaged 356 496 ha and ranged from 4700 to

1 080 500 ha. The subwatersheds averaged 7880

ha and ranged from 96 to 86 500 ha. The 4th- and

6th-code HUCs of the basin assessment area were

characterized by sundry biophysical parameters,

including historic and current vegetation cover

types and structural stages (Hann and others

1997), historic and current terrestrial vegetation

communities (Hann and others 1997), highest

and lowest annual precipitation (available data

from 1989), landform, highest and lowest elevation, and topographic relief.1

The basin assessment area also was characterized

by landform. It contains nine major landforms

ranging from arid grasslands and lowland plains

and valleys, to intermontane basins and breaks, to

steep mountains and glaciated ranges, and a wide

variation in topography, precipitation, and climate

patterns.

Vegetation Cover Types and

Terrestrial Vegetation Communities

This section describes the vegetation classifications used in this analysis and summarizes the

current and historic vegetation character of the

basin assessment area as presented in various

chapters in Quigley and Arbelbide (1997).

Vegetation characterizations of the basin

assessment area—The current vegetation condition of the basin assessment area was characterized by use of 44 vegetation cover types (table 1)

and 24 terrestrial vegetation communities (table 2)

(Hann and others 1997). The vegetation communities are combinations of 41 vegetation cover

types (table 3) and structural stages (see Hann and

others 1997, for descriptions and methods). For

some analyses, vegetation communities better

represent general conditions than do their component cover types.

1 Biophysical data from Intermountain Fire Sciences

Laboratory, USDA Forest Service, P.O. Box 8089,

Missoula, MT 59807.

3
Figure 2—Distribution (hectares) of land ownership and administration in the basin assessment area

(source: ICBEMP, GIS 1-km2 raster data).

Figure 3—Distribution of land by management emphasis, and by unroaded or roaded conditions, in the basin

assessment area (source: ICBEMP, GIS 1-km2 raster data).

4
Table 1—Names of 44 vegetation cover types denoted for

the basin assessment areaa

Cover-type code Vegetation cover-type name

CRB003 Shrub or herb/tree regeneration

CRB005 Alpine tundra

CRB006 Barren

CRB007 Herbaceous wetlands

CRB008 Pacific silver fir/mountain hemlock

CRBS01 Juniper woodlands

CRBS02 Mixed-conifer woodlands

CRBS03 Juniper/sagebrush

CRBS04 Big sagebrush

CRBS05 Shrub wetlands

CRBS06 Agropyron bunchgrass

CRBS07 Native forb

CRBS08 Exotic forbs/annual grass

CRBS09 Grand fir/white fir

CRBS10 White bark pine/alpine larch

CRBS11 Red fir

CRBS12 Cropland/hay/pasture

CRBS13 Fescue-bunchgrass

CRBS19 Urban

CRBS20 Water

SAF205 Mountain hemlock

SAF206 Engelmann spruce/subalpine fir

SAF208 Whitebark pine

SAF210 Interior Douglas-fir

SAF212 Western larch

SAF215 Western white pine

SAF217 Aspen

SAF218 Lodgepole pine

SAF219 Limber pine

SAF227 Western redcedar/western hemlock

SAF233 Oregon white oak

SAF235 Cottonwood/willow

SAF237 Interior ponderosa pine

SAF243 Sierra Nevada mixed conifer

SAF245 Pacific ponderosa pine

SRM104 Antelope bitterbrush/bluebunch wheatgrass

SRM322 Mountain mahogany

SRM402 Mountain big sagebrush

SRM406 Low sage

SRM414 Salt desert shrub

SRM421 Chokecherry/serviceberry/rose a Source: Hann and others (1997). Only 41 cover types are listed here;

3 additional cover types were dropped from the overall list after initial

analyses were conducted for this report. The dropped cover types were

either extremely rare or found to not occur in the basin assessment area

after subsequent study.

5
Table 2—Names and abbreviations of 24 terrestrial vegetation

communities denoted for the basin assessment areaa

Abbreviation Terrestrial vegetation community name

AGRICUL Agricultural

ALPINE Alpine

EXOTICS Exotics

MIXCONES Early seral montane forest

MIXCONLM Late seral montane multilayer

MIXCONLS Late seral montane single layer

MIXCONMS Mid seral montane forest

NFHERBUP Upland herb

NFUPSHRB Upland shrub

PINFORES Early seral ponderosa pine forest

PINFORLM Late seral ponderosa pine forest multilayer

PINFORLS Late seral ponderosa pine forest single layer

PINFORMS Mid seral ponderosa pine forest

RIPHERB Riparian herb

RIPSHRUB Riparian shrub

RIPWDLND Riparian woodland

ROCK Rock/barren

SUBFUPES Early seral subalpine forest

SUBFUPLM Late seral subalpine forest multilayer

SUBFUPLS Late seral subalpine forest single layer

SUBFUPMS Mid seral subalpine forest

URBAN Urban

WATER Water

WOODLDUP Woodland upland

a Source: Hann and others 1997.

6
Table 3—Crosswalk between terrestrial vegetation

communities and vegetation cover types

a

Terrestrial vegetation Vegetation cover-type

community abbreviation code

AGRICUL CRBS12

ALPINE CRB005

EXOTICS CRBS08

MIXCONES CRB003

MIXCONES CRB008

MIXCONES CRBS09

MIXCONES CRBS11

MIXCONES SAF210

MIXCONES SAF212

MIXCONES SAF215

MIXCONES SAF218

MIXCONES SAF227

MIXCONES SAF243

MIXCONLM CRB008

MIXCONLM CRBS09

MIXCONLM CRBS11

MIXCONLM SAF210

MIXCONLM SAF212

MIXCONLM SAF215

MIXCONLM SAF218

MIXCONLM SAF227

MIXCONLM SAF243

MIXCONLS CRBS09

MIXCONLS SAF210

MIXCONLS SAF212

MIXCONLS SAF215

MIXCONLS SAF218

MIXCONLS SAF227

MIXCONLS SAF243

MIXCONMS CRB008

MIXCONMS CRBS09

MIXCONMS CRBS11

MIXCONMS SAF210

MIXCONMS SAF212

MIXCONMS SAF215

MIXCONMS SAF218

MIXCONMS SAF227

MIXCONMS SAF243

NFHERBUP CRBS06

NFHERBUP CRBS07

NFHERBUP CRBS13

NFUPSHRB CRBS04

NFUPSHRB SRM104

NFUPSHRB SRM322

NFUPSHRB SRM402

NFUPSHRB SRM406

7
Table 3—Crosswalk between terrestrial vegetation

communities and vegetation cover typesa (continued)

Terrestrial vegetation Vegetation cover-type

community abbreviation code

NFUPSHRB SRM414

NFUPSHRB SRM421

PINFORES SAF237

PINFORES SAF245

PINFORLM SAF237

PINFORLM SAF245

PINFORLS SAF237

PINFORLS SAF245

PINFORMS SAF237

PINFORMS SAF245

RIPHERB CRB007

RIPSHRUB CRBS05

RIPWDLND SAF217

RIPWDLND SAF235

ROCK CRB006

SUBFUPES CRBS10

SUBFUPES SAF205

SUBFUPES SAF206

SUBFUPES SAF208

SUBFUPLM CRBS10

SUBFUPLM SAF205

SUBFUPLM SAF206

SUBFUPLM SAF208

SUBFUPLS SAF205

SUBFUPLS SAF208

SUBFUPMS CRBS10

SUBFUPMS SAF205

SUBFUPMS SAF206

SUBFUPMS SAF208

URBAN CRBS19

WATER CRBS20

WOODLDUP CRBS01

WOODLDUP CRBS02

WOODLDUP CRBS03

WOODLDUP SAF219

WOODLDUP SAF233

a Terrestrial vegetation communities are a combination of vegetation

cover types and structural stages (not shown here). See table 2 for the

terrestrial vegetation community names associated with the vegetation

community abbreviations and table 1 for vegetation cover-type names

associated with the cover-type codes.

8
Present vegetation conditions—At present, the

five most dominant vegetation cover types of the

basin assessment area (all lands and ownerships)

are big sagebrush (16 percent of the assessment

area), cropland-hayland-pasture (16 percent), interior ponderosa pine (Pinus ponderosa)2 forest

(10 percent), interior Douglas-fir (Pseudotsuga

menziesii) (8 percent), and lodgepole pine (Pinus

contorta) forest (7 percent). The croplandhayland-pasture has been carved mostly from

native grassland types.

Terrestrial vegetation communities are a combination of cover type and structural stage. As

such, the terrestrial vegetation communities show

similar patterns of dominance as those of vegetation cover types and structural stages, but additionally help reveal differences among some of

the general seral forest conditions. The five most

dominant terrestrial vegetation communities of

the basin assessment area (all lands and ownerships) at present are upland shrub (26 percent of

the assessment area), mid seral montane forest

(17 percent), agricultural (16 percent), early seral

montane forest (8 percent), and mid seral ponderosa pine forest (8 percent) (fig. 4).

Change since historic conditions—Hann and

others (1997) present data on change of vegetation cover types and terrestrial vegetation communities since early historic times (about early

1800s). The vegetation cover types having undergone the greatest historic decline in percentage

of original area occupied in the basin assessment

area (that is, less than -50 percent change from

historic area), are western white pine (Pinus

monticola) forest (-96 percent change), whitebark

pine (Pinus albicaulis)/subalpine larch (Larix

lyallii) forest (-95 percent), native forb (-92 percent), Agropyron bunchgrass (-68 percent),

Fescue bunchgrass (-66 percent), cottonwood

(Populus spp.)/willow (Salix spp.) (-64 percent),

and shrub wetlands (-61 percent).3 The vegetation cover types having undergone the greatest

historic increases (that is, > +50 percent change

from historic area) are mountain hemlock (Tsuga

mertensiana) forest (53 percent), Pacific ponderosa pine (80 percent), juniper (Juniperus spp.)/

sagebrush (Artemesia spp.) (163 percent), mountain mahogany (Cercocarpus ledifolius) (406 percent), western redcedar (Thuja plicata) western

hemlock (Tsuga heterophylla) (853 percent),

grand fir (Abies grandis) white fir (Abies

concolor) (965 percent), and Pacific silver fir

(Abies amabilis) mountain hemlock (1,733

percent). Many of the increases in these forest

types have resulted from selective high-grade

logging (for example, of old-growth ponderosa

pine trees) and fire-suppression activities, resulting in major changes in tree dominance and stand

structure (see Hann and others [1997], for data on

absolute changes of each cover type).

The terrestrial vegetation communities decreasing

the most since historic time (that is, less than -50

percent) (fig. 5) include late seral lower montane

(including ponderosa pine) single-layer forest

(-81 percent change), early seral lower montane

forest (-77 percent), upland herb (-67 percent),

and late seral subalpine multilayer forest (-64 percent). Terrestrial vegetation communities most

increasing since historic time (that is, greater than

50 percent) include mid seral lower montane forest (53 percent) and mid seral montane forest (59

percent). Urban areas, exotic communities, and

agricultural areas also have greatly increased, as

they were virtually absent under early historic

conditions. They were carved mostly from native

upland grassland, herbland, and shrubland communities (fig. 6; see Hann and others [1997] for

definitions of urban, exotic, and agricultural communities). Changes in rangeland communities

have been due to increased fire frequency associated with the invasion of exotic grasses, primarily cheatgrass (Bromus tectorum).

2 Authorities for species can be found in the following

references: mosses and nonvascular plants, Schofield

(1980) and Vitt and others (1988); plants and allies,

Hitchcock and others (1969); invertebrates, Furniss and

Carolin (1980); amphibians and reptiles, Collins (1990);

birds, AOU (1983 with supplements); mammals, Jones and

others (1992) and Wilson and Reeder (1993). 3 In addition, Sierra Nevada mixed-conifer forest has undergone significant decline, although this cover type may have

been identified as an artifact within the basin assessment

areas and is not included here in the analyses.

9
Figure 4—Current distribution of terrestrial vegetation communities in the basin assessment area (source:

ICBEMP, GIS 1-km2 raster data).

Figure 5—Percentage of changes of area covered by terrestrial vegetation communities

from historic (early 1800s) to current time, in the basin assessment area: * = not

historically present, lowmont = lower montane, subalp = subalpine (source: ICBEMP,

GIS 1-km2 raster data).

10
Combining terrestrial vegetation communities of

similar seral conditions but different forest types

simplifies the picture (fig. 7). The greatest historic declines have occurred in late seral forests

(-44 percent) and native upland types (including

native grasslands and shrublands, collectively

-38 percent). The greatest historic increases have

occurred in mid seral forests (48 percent) as well

as urban, exotics, and agricultural communities.

Since historic times, there has been a slight increase in overall forest cover (fig. 8) because fire

suppression has encouraged expansion of juniper

woodland and interior Douglas-fir forest into native sagebrush and grassland communities. Of all

forest communities combined, early seral forests

have declined slightly, mid seral forests increased

by about half again, and late seral forests have

decreased by half of their original area (fig. 8).

Separating out these seral stages, we can see how

forest types have differentially changed (fig. 9).

Of the slight overall decline in area of early seral

forests, ponderosa pine forest has declined the

greatest, montane forest has declined somewhat,

and subalpine forest has increased in area (fig.

9a). Of the overall increase in area of mid seral

forests, montane forest and ponderosa pine forest

have greatly increased, while subalpine forest has

remained relatively constant in area (fig. 9b). Of

the overall sharp decline in area of late seral forests, ponderosa pine single-layer forest and subalpine forest multilayer forest have had the

greatest declines; ponderosa pine multilayer forest and montane multilayer forest have declined

somewhat less; and montane single-layer forest

and subalpine single-layer forest have slightly

increased (fig. 9c). We will see later that these

changes have ramifications for associated plant

and animal species.

Figure 6—Historic (early 1800s) and current area of nonforest vegetation communities of the basin assessment

area, showing increase in areas of agricultural and exotic vegetation (source: ICBEMP, GIS 1-km2 raster data).

11
Figure 7—Percentage of changes of area covered by terrestrial vegetation communities,

combined into seral stage categories, from historic (early 1800s) to current time, in the

basin assessment area: * = not historically present (source: ICBEMP, GIS 1-km2 raster

data).

Figure 8—Historic (early 1800s) and current area of forest vegetation communities combined by seral stage, in the basin assessment area (source: ICBEMP,

GIS 1-km2 raster data).

12
Figure 9—Historic (early 1800s) and current area of

individual forest vegetation communities by (A) early,

(B) mid, and (C) late seral stage, in the basin assessment area; lowmont = lower montane, subalp = subalpine (source: ICBEMP, GIS 1-km2 raster data).

13
Methods

Geographic Area Estimates

Estimates of geographic areas are from

ArcInfo®1 geographic information systems (GIS)

1-km2 raster data, provided by the ICBEMP.

Species Data Collection

Data on species distribution and ecology were

gathered from literature and panels of species

experts. Methods for data collection and lists of

experts consulted were presented in Marcot and

others (in prep.).

Further Explorations of Terrestrial

Biodiversity

Global patterns of species richness (number of

species) by taxonomic class were compared with

those of the basin assessment area to determine

differences or similarities. Global richness values

were tallied from the literature, which presented

number of known species and low and high estimated values. Values of species richness from

the basin assessment area were tallied as single

values

for number of known species and number of estimated species (Marcot and others 1997). The

poorest known taxonomic classes are those with

the lowest proportion of estimated species that

have been scientifically verified from the assessment area. These were compared with global patterns. Implications of terrestrial biodiversity

patterns for management are identified based

on similarities of these patterns.

Assessment of Macroecology of

Individual Species and Species

Groups

Species database—A total of 1,347 individual

terrestrial species and 93 species groups of

plants and animals were represented in a speciesenvironment relations (SER) database (Marcot

1997; Marcot and others 1997). These are not all

the individual species or species groups that occur

in the basin assessment area but includes rare or

potentially rare plants, selected invertebrates, and

all vertebrates.

The SER database includes descriptions of (a) key

environmental correlates of species, which consist

of macrohabitats (vegetation cover types and

structural stages) and other microhabitat and environmental factors that affect the presence, distribution, and realized fitness of species; (b) KEFs

of species; (c) ecological status, including degree

of endemism and geographic distribution of species; (d) legal listing status of species by various

Federal and State agencies; and other information.

Species groups included lichens, bryophytes, vascular plants, bacteria, protozoa, rotifers, and nematodes. Species of lichens and bryophytes were

grouped according to common use of substrates or

soil conditions. Species of vascular plants were

grouped based on various criteria, including genera difficult to identify, taxonomic uncertainty,

and taxa for which knowledge and experts were

available. Some vascular plant groups likely are

either underrepresented or not represented in

some geographic areas. Thus, although we summarize number of vascular plant groups, we do

not include them in subsequent ecological analyses. Bacteria, protozoa, rotifers, and nematodes

were each grouped according to their presence

and ecological function in soils.

1 The use of trade or firm names in this publication is for

reader information and does not imply endorsement by the

U.S. Department of Agriculture of any product or service.

Chapter 3

14
In the SER database, information on rare or potentially rare plants also included life form, categories of population trend (as denoted in state

Natural Heritage databases), dispersal mode, pollinators, indicators of specific soil or environmental conditions, and occurrence by state, general geographic area, substrate, and local topographic condition (azimuth, relief, and slope). Information on invertebrates also included dispersal

mode. Information on vertebrates also included

percentage of overall distributional range of each

species occurring within the basin assessment

area, and the breeding, seasonal occurrence, and

abundance status of birds.

The SER database was queried to produce ecological profiles of species by taxonomic group,

habitat condition, ecological function, ecological

status and geographic occurrence, legal status,

and other attributes. Because few studies have

been conducted on most of the basin species, the

SER database is incomplete and consists largely

of categorical information based on the judgments and experiences of experts (see Marcot

and others [1997] and Marcot [1997] for caveats

on use and interpretation of the SER database

information).

Plants and allies—Occurrence and autecology

of fungi, lichens, bryophytes, and vascular plants

of the basin assessment area were summarized in

Marcot and others (1997) and will be detailed in

several other publications. In this report, number of taxa and species groups of plants and allies

were tallied by life-form category, population

trend class, geographic occurrence, substrate,

local topographic condition, dispersal mode, type

of pollinator, and type of indicator function. Lifeform categories of plants followed the classification of Raunkaier (1934). Information in the SER

database on plant dispersal modes was entered as

comments taken from experts; the comments

were then coded into categories, as discussed in

“Results.” Trends of vascular plant species, and

of lichen, bryophyte, and vascular plant groups

were denoted in four categories (unknown, decreasing, stable, and increasing) by the Plant Task

Group of the Science Integration Team. In this

document, only scientific names are used for

plants, as common names have not been

standardized.

Invertebrates—Ecology of invertebrates of

the basin assessment area was noted in Marcot

and others (1997), summarized in Niwa and

Sandquist (in prep.), and will be detailed by functional group in several other publications. In this

report, dispersal modes of example invertebrate

species of the basin assessment area were categorized and cross-tabulated with KEFs of species.

This helped determine the role of dispersal ecology in maintaining ecological functions of the

invertebrate fauna. Examples of invertebrate species and micro-organism groups dispersing by

phoresis (by using another organism as a dispersal host) were listed, illustrating how species

interactions have evolved that potentially influence viability and distribution of some invertebrates and micro-organisms.

Vertebrates—Ecology of vertebrates of the basin

assessment area was summarized in Marcot and

others (1997) and will be detailed by selected

species groups in other publications. In this report,

patterns of taxonomic diversity were explored by

tallying number of species, genera, and families

through the basin assessment area and by vegetation cover types and structural stages.

Key ecological functions of species were further

explored by cross-tabulating number of vertebrate

species by ecological functions and by terrestrial

vegetation communities. This resulted in “speciesfunction profiles” that were used to determine

which vegetation communities had the greatest

variation in KEFs of species, and which ecological function categories varied the most over vegetation communities.

Species function profiles were developed for all

vertebrates combined by tallying the number of

species with selected specific KEFs, by terrestrial

vegetation community. Greater numbers of species with the same KEF in a specific community

were interpreted as greater redundancy of that

function. Similarly, Huston (1994:3) referred to

species with the same KEFs as “functional types

of organisms,” and redundancy of species with a

particular KEF as “functionally analogous species.” Franklin and others (1989:93) noted that

“forest ecosystems which have redundancy in

structure and function are more likely to be able

15
to absorb stresses, including species losses, without catastrophic damage. . . .” Silver and others

(1996:17) similarly concluded that “functional

diversity, and not just species richness, is important in maintaining the integrity of nutrient and

energy fluxes,” and that high species richness

affords alternative and redundant pathways for the

flow of resources in an ecosystem. Our species

function profiles helped (1) identify communities

with the greatest (and the least) redundancy in

ecological functions of species, and (2) determine

variation in redundancy of each KEF among

communities.

Total functional diversity of all vertebrate species

was calculated by multiplying the number of

KEFs of vertebrate species in each terrestrial vegetation community by the average number of vertebrate species performing each function (after

Huston 1994:4ff). Species function profiles and

total species functional diversity were not calculated for rare or potentially rare plants and allies,

nor for invertebrates, because (a) scientific

knowledge and our databases of these taxa are incomplete, and (b) most of these species likely respond to microhabitats, microclimates, and substrates at far too fine a resolution than afforded

by the available broad-scale data in this study.

Broad-scale ecological distribution of birds was

assessed by cross-tabulating number of bird species by abundance, breeding status, migration

status, and U.S. state, and relating to general geographic location. Five abundance categories and

five breeding or migration status categories were

denoted for birds (appendix A). Bird species

abundance, breeding status, migration status, and

occurrence by U.S. state were examined for patterns of local and regional endemics (see Marcot

and others [1997] for definitions of endemism

classes used with plants and animals).

We explored potential effects of regional climate

change2 on vertebrates. We first listed species

associated with vegetation communities that may

undergo major declines in distribution and area

under climate-change scenarios. We then tallied

the number of vegetation cover types used by

each species and rank-ordered the species in increasing number of cover types used. We identified species at risk as those that use <20 percent

(and also those species at greater risk that use

<10 percent) of the total number of vegetation

cover types (N = 44). These are species closely

associated with the primary vegetation community of interest and are thus at greatest risk

of decline or extirpation from climate-induced

changes of that community.

Present and Past Terrestrial

Characterization of the Basin

Assessment Area

Present character—The Science Integration

Team described broad-scale characteristics of

the basin assessment area in terms of vegetation cover types and structural stages, geology,

weather, land form, and other attributes, by

1-km2 cells, 4th-code HUCs, and 6th-code HUCs

(chapters in Quigley and Arbelbide 1997).

Historic character—We used results of reconstructions of early historic (early 1800s) vegetation conditions and changes to current conditions,

as provided in Hann and others (1997).

Prehistoric character—In this report, we also

summarize aspects of paleoclimates and paleoecology of the basin assessment area by comparing reports of prehistoric climates and biota with

those of the present. We compared diversity of

vertebrate Tertiary paleofaunas with that of modern faunas in eastern Oregon, at order, family,

genus, and species levels of taxonomy and by

geologic period, and drew conclusions on qualitative changes in environments, turnover of various

taxa, and changes in overall terrestrial communities. Data are lacking on prehistoric rates of species extinction and speciation in the basin assessment area.

Assessment of Ecological Integrity

of Terrestrial Species and

Communities

Ecological integrity refers to the degree to which

native taxa and their ecological functions persist

under human land management (for example,

Majer and Beeston 1996). Ecological integrity

was assumed by the Science Integration Team to

be a general purpose for managing ecosystems.

Ecological integrity reflects the degree to which

all ecological components and their interactions

2 Personal communication. Sue Ferguson, Climate model

data, Forestry Sciences Laboratory, Seattle WA 98105-6497.

16
are present and functioning. We assumed that the

six goals of managing for ecological integrity in

land use planning, as discussed by Haynes and

others (1996), provide important benchmarks

against which to measure progress. Three of those

goals dealt explicitly with terrestrial ecological

conditions: maintaining species viability, maintaining long-term evolutionary potential of species, and managing for multiple ecological domains and evolutionary timeframes. We assumed

that those three ecological goals could be interpreted in terms of species management objectives

by using spatially referenced information on terrestrial species ranges and habitat and environmental conditions. In this way, conditions in 6thcode HUCs were mapped in a GIS (ArcInfo), for

each component of terrestrial ecological integrity.

Specifically, we followed these steps:

1. Reiterate the six goals for ecological integrity

as identified by Haynes and others (1996) and

identify those pertinent to terrestrial ecology

(goals 1-3).

2. Under each ecological integrity goal, identify

components pertinent to terrestrial ecology, particularly those that can be mapped and quantified.

Denote these components under each of the three

goals pertinent to terrestrial ecological conditions

(as discussed in Marcot and others 1997).

3. For each integrity component, identify corresponding species by querying the SER database.

Then identify the set of GIS map themes, including distributions of species, to represent the

component.

4. For each integrity component and its set of GIS

map themes, quantify conditions for each 4th code

HUC in the basin assessment area. Use these data

to build a database in which rows are individual

4th code HUCs, columns are the integrity components, and cells are filled in based on counts,

percentages of each HUC covered, or other measures pertinent to each component.

5. Produce a map for each integrity component

showing, by 4th code HUC, the outcome of

combining all relevant GIS themes as defined in

step 4.

6. Merge the database from step 4 with similar

data on 4th code HUCs into an overall database

expressing combined conditions for all terrestrial

ecology components of ecological integrity.

Overall results were expressed in three maps

depicting current average conditions for each of

the three ecological integrity goals pertaining to

terrestrial species and ecosystems. These maps

are intended to provide a broad-scale overview of

the basin assessment area that may be useful in

planning policy. They are not scaled for site-level

or midscale management use.

17
Results

Further Explorations of Terrestrial

Biodiversity

Marcot and others (1997) present a preliminary

summary of biodiversity patterns of species

among taxonomic groups within the basin assessment area. In this section, we expand on those

findings.

First, how does biodiversity within the basin assessment area compare with global patterns? Estimated global biodiversity suggests that most species are insects (fig. 10, Wilson 1988). Next in

estimated global abundance are algae (not tallied

for the current work), bacteria, fungi, nematodes

(roundworms), spiders and mites, and other taxa.

The least abundant taxa globally are vertebrates

(fig. 10). Estimates of diversity in the basin

assessment area suggest that most species are

bacteria, followed by fungi, macroinvertebrates

(insects), plants and allies (lichens and bryophytes), and other species. The least abundant in

the basin assessment area are soil nematodes1 and

vertebrates (fig. 11).2 Overall patterns indicate

that, both globally and in the basin assessment

area, estimated species diversity (number of species) of invertebrates and micro-organisms rank

highest and vertebrates lowest.

On comparing known with estimated species

diversity, it is obvious that the largest gap in

scientific knowledge (less than half of estimated

numbers of species actually known) globally

occurs with meso- and micro-organisms. The

best known taxa are vertebrates. In the basin

assessment area, the least known taxa probably

include most of the micro-organisms, as well as

insects, fungi, and mollusks. Only 14 percent of

the estimated number of insect species in the

basin assessment area have actually been reported

from the basin assessment area. The percentages

for fungi (33 percent) and mollusks (49 percent)3

are a little greater. (Percentage values presented

here are approximations.) Globally, the percentage of estimated number of species (from the

upper estimate values) that actually are known are

0.13 percent of bacteria, 0.95 percent of insects,

1 percent of viruses, 5 percent of fungi, 8 percent

of spiders and mites, and 35 percent of mollusks.

Thus, on a percentage basis, these taxa are better

known in the basin assessment area than they are

globally, although much basic work on species

inventory and taxonomic studies remains to be

done.

The best known taxa (at least half of estimated

number of species actually known) in the basin

assessment area probably are vertebrates and

lichens (all or nearly all likely occurring species

have probably been discovered and catalogued in

the basin assessment area), followed by vascular

plants (99 percent or perhaps a bit less) and bryophytes (94 percent). Globally, the best known taxa

are vertebrates (90 percent) and vascular plants

(50 percent). Most of the biodiversity patterns of

1 That nematodes rank high in global biodiversity and low in

the basin assessment area likely reflects how poorly known

they are in the basin assessment area rather than their true

abundance. All the values of known and estimated species

biodiversity are subject to change as new studies are

initiated.

2 This comparison between global and basin assessment

area biodiversity is not strictly at parity, because data on all

taxonomic groups as divided for the basin assessment area

analysis were not available at a global level.

3 No figures for these percentages are available for

micro-organisms in the basin assessment area. So little work

has been done on these taxa that virtually all are unstudied

and empirically undocumented in the basin assessment area.

Chapter 4

18
Figure 10—Diversity of terrestrial species globally, showing known, low estimates, and

high estimates of number of species by taxonomic class, sorted in decreasing order of

high estimates (source: Marcot and others 1997). (Note log scale.)

Figure 11—Diversity of terrestrial species and fish in the basin assessment area,

showing known and estimated number of species by taxonomic class, sorted in

decreasing order of estimated number. (Note log scale.)

19
species numbers by major taxonomic groups in

the basin assessment area follow global patterns.

Some groups are better known in the basin assessment area because the global estimates are

weighted heavily toward largely unstudied

tropical ecosystems.

Macroecology of Individual

Species and Species Groups

Plants and Allies

Number of species—In the basin assessment

area, some 18,946 species of plants and allies

are estimated to exist (Marcot and others 1997).

This figure extrapolates for species not yet discovered or catalogued and includes projections of

about 9,000 species of macrofungi, 736 lichens,

860 bryophytes, and 8,350 vascular plants. At

present, only 12,797 species of plants and allies

(68 percent of all estimated species) are actually

known from the basin assessment area, including

about 3,000 macrofungi (33 percent of all estimated macrofungi species), 736 lichens (100 percent of all estimated lichens), 811 bryophytes (94

percent of all estimated bryophytes), and 8,250

vascular plants (99 percent of all estimated vascular plants). An additional large set of microfungi,

including mycorrhizae, likely occur but are

largely unstudied and are not treated here.

The SER database lists 596 individual taxa

(mostly species with selected subspecies or varieties) and 82 species groups of plants and allies.

These include 394 species of macrofungi; 2 species and 39 groups of lichens; no species and 11

groups of bryophytes; and 200 taxa and 32 groups

of vascular plants (one part of the SER database

also lists 920 vascular plant taxa, but information

on environmental correlates and ecological functions are not available for all of these taxa.) The

596 taxa and 82 groups mostly represent rare or

potentially rare but largely unstudied taxa and

communities; the vascular plants included are

mostly perennial forbs or nonwoody perennials.

Thus, the analyses that follow pertain only to this

select subset of rare or potentially rare plants of

the basin assessment area and do not represent

conditions of the full flora.

Species and groups by geographic area—

Presence by general geographic area within the

basin assessment area was denoted for lichen

groups, vascular plant groups, and vascular plant

species. Numbers of lichen groups and vascular

plant groups were relatively evenly distributed

throughout most of the geographic areas, with

fewer in Wyoming and Nevada probably because

of the small total area these states occupy within

the basin assessment area. Exceptions also include the Columbia Plateau, which is entirely

contained within the basin assessment area but

which had only two lichen groups represented,

and the Basin and Range area (northern Great

Basin), which extends well beyond the basin assessment area and had only one vascular plant

group. The few numbers of these species groups

may be an artifact of the grouping process.

Numbers of the rare and potentially rare vascular

plant species considered in this assessment were

less evenly distributed among general geographic

areas (fig. 12) than were lichen groups and vascular plant groups. Geographic areas with more

species (>25 species) included Basin and Range,

Columbia Plateau, Blue Mountains, northern

Idaho, and the southern part of the east side of

the Cascade Range. Areas with fewest species

(<10 species) included the small portion of

Wyoming within the basin assessment area,

southern Idaho, and Okanogan Highlands,

although the last two of these areas were not

reduced in number of species groups.

Life forms—Life forms of most of the rare or

potentially rare fungi were mushrooms (59 percent of all 394 macrofungi taxa included in the

SER database) and truffles (31 percent), with the

rest puffballs (7 percent), polypores (2 percent),

and resupinate (1 percent) (fig. 13; see definitions in table 4).

Life forms of most of the rare or potentially rare

vascular plants were hemicryptophytes (36 percent of all 280 vascular plant taxa included in the

SER database) and cryptophytes (30 percent),

with the rest therophytes (16 percent), chamaephytes (15 percent), and phanerophytes (4 percent) (see definitions in table 4). Plants of most

of these life-form categories are widely distributed among vegetation cover types of the basin

assessment area. The hemicryptophytes (102 species) are found in 27 vegetation cover types, most

20
Figure 12—Number of rare or potentially rare vascular plant species by geographic area

within the basin assessment area.

Figure 13—Diversity of macrofungi in the basin assessment area by life form (see table 4

for descriptions of life forms).

21
Table 4—Classes and definitions of fungi and vascular plant life formsa

Class Definition

Fungi:

Mushroom Fungi with any macrofruiting body, typically those with cap and stipe

morphology

Resupinate Upside down or recurved growth form

Truffle Fungi that are hypogeous (undergound), rare, odoriferous, and edible, of

Ascomycetes

Puffball Fungi of Gastromycetes of the order Lycoperdales (true puffballs and stalked

puffballs); does not include Hymenogastrales (false puffballs)

Polypore Fungi in the family Polyporaceae

Vascular plants:

Phanerophyte Species with perennating buds or shoot apices on aerial shoots. Group

includes most woody shrubs or trees

Chamaephyte Species that hold their perennating buds and shoot apices close to the

ground. Group includes semiwoody and herbaceous species that persist

aboveground throughout the year (though not necessarily in a

physiologically active state). Group includes bunch grasses and cushion

plants

Hemicryptotphyte Species with perennating buds at ground level, generally protected by snow

or organic debris, with all plant parts aboveground dying back at the end of

the active growing season. Stolons may or may not be present. Group

includes rosette plants (for example, Taraxacum), partial rosette plants (for

example, Achillea millefolium), and stoloniferous species (for example, Rubus)

Cryptophyte Species bear their perennating buds below ground level or submerged in

water. Group includes rhizomatous and bulb-forming species (for example,

Allium), and aquatic species such as Alisma and Nuphar

Therophyte Annual species (for example, Bromus tectorum, Stephanomeria malheurensis).

Embryonic buds are protected by a seed coat

a Fungi definitions from W. Owen. Vascular plant definitions from L. Croft based on Raunkier (1934); also see

Daubenmire (1974).

22
principally in big sagebrush (26 species), low

sage (14 species), mountain big sagebrush (13

species), fescue-bunchgrass (12 species), and

alpine tundra (11 species), with the remaining

26 species distributed among 22 other vegetation

cover types. The cryptophytes (84 species) are

found in 22 vegetation cover types, most principally in fescue-bunchgrass (20 species),

Agropyron bunchgrass (14 species), interior

ponderosa pine forest (13 species), and juniper/

sagebrush (10 species), with the remaining 27

species distributed among 18 vegetation cover

types. The therophytes (45 species) are found in

21 vegetation cover types, most principally in big

sagebrush (10 species), juniper sagebrush (9 species), interior ponderosa pine (9 species), and

interior Douglas-fir (8 species), with the remaining 9 species distributed among 17 other vegetation cover types. The chamaephytes (42 species)

are found in 21 vegetation cover types, most

principally in low sage (13 species), big sagebrush (8 species), and mountain big sagebrush

(5 species), with the remaining 16 species distributed among 18 other vegetation cover types.

Finally, the phanerophytes (10 species) are found

in seven vegetation cover types, most principally

in Agropyron bunchgrass (two species) and low

sage (two species), with the remaining six

species distributed among three other vegetation

cover types. These are very general patterns;

information on finer classes of habitat types

would reveal more useful botanical patterns.

Number of rare or potentially rare vascular plant

species among life-form categories do not show

a clear trend at the broad scale in relation to general topographic position, soil texture, soil depth,

soil moisture regime, or soil temperature categories, although correlations would be expected at

finer scales of resolution. This is not surprising,

given the very microsite- and substrate-specific

nature of most of these rare or potentially rare

plants. Based on the initial assignments of plants

to life-form categories, hemicryptophytes might

occur more frequently in calcareous substrates

than do other life forms; they also might occur

most commonly in alkaline soil conditions. This

may represent an ecological pattern, or merely

reflect the dominance of this life form in the rare

plant flora. These patterns need verification and

further ecological study.

In general, alkaline or calcareous soils are typically rich in edaphic endemics worldwide. Most

plant species are unable to overcome the physiological challenges imposed by such soils, including Ca++/Mg++ imbalances and the general low

availability of nitrogen, phosphorus, and potassium nutrients in high pH situations because of

specific solubilities. Alkaline or calcareous sites

tend to be azonal; that is, represented by an intrusion into a matrix of some other parent material (for example, granite or sandstone). Those

plants that thrive in such azonal settings with few

other species capable of surviving the local conditions often have an opportunity to escape competition. Azonal soils almost always have unique,

if not rare, flora. Examples include serpentine or

ultramafic conditions in northwestern California

and southwestern Oregon, volcanic ash in eastern

Oregon and western Idaho, and hyperacidic mine

tailings, hot springs, and sandy substrates elsewhere. Specialization of rare or endemic plants to

azonal conditions can be considered an extreme

form of habitat segregation.

Trends of vascular plant species—Of 299 rare

or potentially rare vascular plant species, 143 species (48 percent) had unknown or unreported

trends, 84 species (28 percent) were coded as decreasing, and 71 species (24 percent) as stable.

One species (<1 percent) might be increasing at

least in some areas. Among the decreasers were

various species of Allium, Aster, Astragalus,

Botrychium, Calochortus, Cypripedium,

Eriogonum, Haplopappus, Howellia, Lepidium,

Mentzelia, Mimulus, Penstemon, Ranunculus,

Silene, Thelypodium, Trifolium, and other genera.

Among the stable species were various other

species of Allium, Astragalus, Botrychium,

Castilleja, Claytonia, Erigeron, Eriogonum,

Leptodactylon, Mentzelia, Mimulus, Penstemon,

Trifolium, and other genera. The sole potential

increaser was an Astragalus species. Species with

unknown or unreported trends spanned an even

greater number of genera.

Trends of lichen, bryophyte, and vascular

plant groups—Trends were denoted for lichen

groups, bryophyte groups, and vascular plant

groups (fig. 14). Trends of most of the lichen

groups were decreasing (41 percent of all lichen

groups) or stable 36 percent), with fewer groups

having unknown or unreported trends (18 percent)

or increasing trends (5 percent). Trends of

23
bryophyte groups were largely unknown or unreported (83 percent of all bryophyte groups),

with fewer decreasing (17 percent); none was

stable or increasing. Trends of vascular plant

groups were mostly unknown or unreported (44

percent of all vascular plant groups), with fewer

groups stable (33 percent) or decreasing (22 percent); none was increasing.

The only two lichen groups noted as having increasing trends were excess nitrogen-indicator

lichens and urban pollution-tolerant lichens.

The 28 stable groups included 16 lichen groups

in various aspen, calcareous, fencepost, leaf, rock,

seepage, ledge, and tundra conditions and substrates, and 12 vascular plant groups consisting

of 2 Allium groups, 5 Mimulus groups, and 5

Penstemon groups.

The 28 decreaser groups included 18 lichen, 2

bryophyte, and 8 vascular plant groups in various

arid, riparian, and forest conditions. The two

decreaser bryophyte groups included the decayed

wood bryophyte group and the epiphytic bryophyte group.

The 24 lichen groups with unknown or unreported

trends included 3 lichen groups of bog, nitrogen

fixation, and pioneer soil stabilizer conditions; 9

bryophyte groups of aquatic, humid duff, peatland, rock, and soil conditions; and 12 vascular

plant groups, all Carex groups mostly of meadows association, with a few of peatland and forest conditions.

Lichen, bryophyte, and vascular plant species

groups by substrate—Lichen, bryophyte, and

plant groups included in the SER database were

denoted with orientations to specific nonvegetation substrates and vegetation substrates.

Patterns of species groups among nonvegetation

substrates were as follows: lichen groups were

more numerous on basalt rock, other rock, and

welded tuff; bryophyte groups were fairly evenly

distributed among all substrates but were absent

on pumice and welded tuff (fig. 15). Among all

taxa, wet and dry soil substrates, and basalt and

other rock substrates were used by the most species groups, and pumice and alkali soils were

used by the fewest species groups (fig. 15). The

composition of species groups, however, differed

among these sets. For example, the sole lichen

Figure 14—Number of lichen, bryophyte, and vascular plant species groups in the basin

assessment area, by trend category.

24
group found associated with pumice substrates

was the calcareous indicator lichens group. This

group also is found on five other nonvegetation

substrates. The two lichen groups—the pioneer

soil stabilizer lichen group and the soil lichen

group—associated with tuffaceous ash each also

are found in three other nonvegetation substrates

(dry, wet, and alkali soils).

Patterns of species groups among vegetation substrates were as follows: lichen groups were most

numerous as epiphytic conditions and were absent in peatland, fen, and bog substrates (although

individual lichen species may occur in such conditions); bryophyte groups were most numerous

in aquatic submerged substrates but occurred in

all but musicolous substrates; and vascular plant

groups were most numerous in peatland, fen, and

bog substrates and also occurred in humus and

duff substrates but were not identified for the

other vegetation substrates (although individual

species occur on such substrates) (fig. 16).

Patterns of plant endemism—There are many

examples of endemic plants in the basin assessment area. Some examples are Idaho goldenweed

(Haplopappus aberrans) in the Boise River drainage, Chaenactis evermannii Greene in central

Idaho, and C. nevii Gray in the John Day area.

Not all endemic plants are rare, and not all rare

plants are endemics. A large proportion of the

rare or potentially rare plants considered in this

assessment, however, are locally endemic or

regionally endemic (156 taxa or 78 percent of

200 vascular plant species in the SER database

with information on endemism status). We defined local endemic as a plant with populations

restricted to a very small area such as one portion

of a mountain range or one canyon. Local endemics also may be specialized on highly restricted

habitats. The entire range may lie within the basin

assessment area. A regional endemic is a plant

with populations that inhabit a larger geographic

area than that of local endemics such as southeast

Oregon or the Palouse. The range of regional endemics may extend beyond the basin assessment

area boundaries. Regionally endemic plants also

may be closely associated with specific habitats.

In general, about a third of all vascular plants

(including common and rare species) are endemic

at least regionally.

Figure 15—Number of lichen, bryophyte, and vascular plant species groups by soil and

rock (nonvegetation) substrate condition in the basin assessment area.

25
Gentry (1986) tallied endemic plants (mostly

based on candidate taxa for Federal listing, similar to our consideration of plants of conservation

concern) by U.S. state. We compare our tallies

with his in table 5. His definition of endemic

taxa included plant species with distributions of

50 000 km2 or less, which corresponds to our

categories of local and regional endemics. His

tallies included entire states, whereas ours included only portions of states within the basin assessment area. We counted more endemic plants

in Idaho and Montana than he did, however;

whereas his figures for Oregon, Washington,

Nevada, and Wyoming were greater than ours.

(We did not characterize plant endemism in the

Utah portion of the basin assessment area.) Only

a small portion of Nevada and Wyoming are within the basin assessment area, so these differences

are not surprising. But we were surprised to see

our greater tallies for Idaho and Montana, considering particularly that only the western portion

of Montana (west of the Continental Divide) was

included in the basin assessment area, and only a

portion of the entire flora of the basin assessment

area is included in the SER database. Our greater

tallies of endemic plants in Idaho and Montana

possibly reflect more current data from state

Natural Heritage program databases.

Gentry (1986) suggested two kinds of endemics.

The first kind is paleoendemics, which are species that were formerly more widespread but now

have restricted distributions because of climatic

changes and natural or human-caused reduction

of favorable habitats. Some paleoendemics may

be archaic taxa with primitive traits; some may

be close to natural extinction. Pielou (1991:240)

notes that the present alpine flora of the western

mountains contains a mix of paleoendemic plant

species, some presumably derived from Pleistocene refugia south of the Cordilleran ice sheet

and others from northern Beringian tundra. She

also noted that the nearest unglaciated drainage

basin to the Cordilleran ice of the Pleistocene was

that of the Columbia River, which must have

been the refuge for many paleoendemic species.

The second kind of endemic is neoendemics,

which are species that have recently evolved and

have restricted ranges because they have not yet

had time to spread over a larger geographic area.

Examples of neoendemics might include species

Figure 16—Number of lichen, bryophyte, and vascular plant species groups by vegetation

substrate condition in the basin assessment area.

26
that have recently arisen on serpentine and ultramafic substrates. Most species of Astragalus in

the basin assessment area likely are neoendemics.

Allium aaseae (Ownbey) is a “classic”

neoendemic.

Kruckeberg and Rabinowitz (1985) suggested

that paleoendemics tend to have more than

one disjunct population, whereas endemics

confined to a single population can be either

paleoendemics or neoendemics. This contention

is untested for the regional or local endemic

plants of the basin assessment area. Of course,

one cannot assert the converse; plants with more

than one disjunct population cannot be assumed

to be a paleoendemic, any more than plants with

a single population are endemics of either type.

In general, paleontology explains much about

current plant species distributions, especially

with respect to historic climates. Some plants of

northern Idaho occur because of the Clearwater

Maritime refugium found there. Southern Idaho

and the portion of Wyoming in the basin assessment area are geologically young, and the few

endemics found there are classic neoendemics

of species-rich genera (for example, Astragalus).

Also, climate fluctuations over the last several

million years in southern Idaho have been broad,

thereby making it difficult for either mesophytes

or xerophytes to persist over the entire period

(Tausch and others 1993).

Overall, genetic data are needed to determine

which other vascular plant taxa within the basin

assessment area are paleoendemic or neoendemic.

Chemical, physical, and biological properties of

soils often constrain endemic taxa (Kruckeberg

and Rabinowitz 1985). In some cases, rarity or

range-restriction of a coevolved symbiont may

account for endemism of plant taxa, particularly

with a rare obligate pollinator but, again, this

needs empirical study in the basin assessment

area.

We explored the question of environmental determinants of endemic plants by comparing 13 biophysical factors between the 156 plant taxa from

the SER database that were denoted as local or

regional endemics to the remaining 44 plant taxa

in the database denoted as neither local nor regional endemics. This analysis is pertinent only

Table 5—Tallies of locally and regionally endemic plant taxa in

U.S. states of the basin assessment area, compared with estimates

of endemic plant species from Gentry (1986:155)a

This study

Endemics

Local Regional as listed by

State endemics endemics Total Gentry (1986)

Oregon 47 30 77 109

Washington 24 13 37 49

Idaho 29 22 51 37

Montana 6 7 13 6

Nevada 2 1 3 19

Wyoming 1 3 4 90

Entire basin

assessment

area 93 63 156 —

— = no data a We defined local endemic as plants with populations restricted to a very small area

such as 1 portion of a mountain range or 1 canyon, and regional endemic as plants

with populations that inhabit a larger geographic area such as southeast Oregon or

the Palouse. Gentry defined endemic as plant species with distributions of 50 000 km2

sor less, which corresponds to our categories of both local and regional endemics.

27
to rare or potentially rare plant taxa and does not

describe more common or widespread plant species. The biophysical factors included in this analysis were maximum elevation, minimum elevation, aspect, slope angle, slope position, geographic area, geology, landform, soil pH, soil texture, soil depth, soil moisture, and soil temperature. These factors were coded for each species in

the SER database as categorical values. We used

a log-likelihood ratio contingency test to analyze

the null hypotheses that local and regional endemics do not differ from nonendemics, in number of

species distributed among categories of each of

these biophysical factors.

Results are presented in table 6. Only two biophysical factors—slope angle and slope position—were significantly different (P < 0.05)

between endemic and nonendemic rare plants.

Six other factors were marginal in significance

(0.05 < P < 0.10): maximum elevation, aspect,

geology, landform, soil pH, and soil moisture.

Given the broad scale of the basin assessment

area and coarse resolution of the categorical data

denoted in the SER database, the marginally significant factors might be seen as working hypotheses of biophysical conditions that influence numbers of rare endemic plants. Finer grain and more

quantitative data, as well as species-specific analyses, are needed to further unravel the factors

inducing plant endemism in the basin assessment

area. Also needed are further studies to explain

the exclusion of endemics from more common

habitats (Kruckeberg and Rabinowitz 1985).

Plant dispersal modes—Dispersal (dissemination) modes of all rare or potentially rare plants

of the basin assessment area are not thoroughly

studied, but those known were denoted in the

SER database. Seven general dispersal mode

categories were identified: gravity, wind

(anemochory), water, insects, vertebrates

(zoochory), growth or reproduction, and unknown. The vertebrates dispersal category was

further divided into dispersal by birds, small

mammals, large mammals, and not specified.

The growth or reproduction dispersal category

was divided into dispersal by dehiscence, bulbett

scattering, spore dispersal, stolon, rhizome, vegetation growth, seed dispersal, and turion. Dispersal mode was denoted for lichen, bryophyte, and

vascular plant groups, and for the rare or potentially rare species of fungi and vascular plants.

Any one group or species could have been coded

as having one or more dispersal modes.

Gravity plays the major role in dispersal of nearly

all the lichen groups, with one other group (aquatic rock lichens) dispersing by water and by vertebrates (birds) (fig. 17). Dispersal modes of bryophyte groups were evenly split between water and

wind.

About two-thirds (69 percent) of the 394 rare or

potentially rare fungi species included in the

assessment disperse by wind, and the remainder

disperse by vertebrates (that is, mycophagy) (fig.

18). Mycophagy, or the ingesting and distributing

of belowground fungal bodies (truffles) including

spores, is an important ecological role of some

large and small vertebrates in several forest types

throughout the inland West. Vertebrates engaged

in mycophagy and dispersal of fungi include

Rocky Mountain elk (Cervus elaphus nelsonii),

western red-backed vole (Clethrionomys

californicus), southern red-backed vole (C.

gapperi), northern flying squirrel (Glaucomys

sabrinus), least chipmunk (Tamias minimus),

Douglas squirrel (Tamiasciurus douglasi),

American pika (Ochotona princeps), and

Columbian mouse (Peromyscus keenii).

Dispersal modes of vascular plant groups included all seven major dispersal categories. Most

vascular plant groups, however, disperse mainly

by gravity, wind, water, and vertebrates (fig. 17).

The rare or potentially rare vascular plant species

disperse (in order of decreasing number of species per dispersal category) by gravity (27 percent

of species), wind (26 percent), growth or reproduction (21 percent), vertebrates (13 percent),

water (7 percent), and insects (2 percent). How

the remainder disperse (4 percent) is unknown

(fig. 18). The 55 vascular plant species that disperse by vertebrates more or less evenly disperse

by birds and small mammals, with fewer species

dispersing by large mammals, and a third not

specified (fig. 19a). Over half of the 89 vascular

plant species that disperse by growth or reproduction do so by seed set; the remainder disperse

by many other vegetative and reproductive means

(fig. 19b). It must be remembered, though, that

these patterns pertain only to the set of rare or

potentially rare plants considered in this assessment, and that patterns of the full flora—

28
Table 6—Results of log-likelihood ratio contingency tests (G statistic) of biophysical factors and endemic plantsa

Biophysical factor Value of G df P

Maximum elevation: 6.678 3 0.05 < P < 0.10 +

<1219 meters

1219 to 1828 meters

1829 to 2438 meters

2439+ meters +

Minimum elevation: 1.443 1 0.10 < P < 0.25

<1219 meters

1219+ meters

(Categories combined because of

small sample sizes)

Aspect: 7.781 4 0.05 < P < 0.10 +

North

South

East

West

Flat

Slope angle: 7.929 3 0.025 < P < 0.05 *

Flat (0 to 10 percent)

Gentle (11 to 30 percent)

Steep (31 to 50 percent)

Very steep (>50 percent)

Slope position: 7.155 2 0.025 < P < 0.05 *

Lower slope

Middle slope

Upper slope

Geographic area: 11.743 7 0.10 < P < 0.25

Basin and range

Blue Mountains

Columbia basin

East Cascade Range north

East Cascade Range south

Montana

Northern Idaho

Southern Idaho

(Other states omitted because of

small sample sizes)

Geology: 16.178 10 0.05 < P < 0.10 +

Basalt

Calcareous

Clay

Glacial

Granite

Playa

29
Table 6—Results of log-likelihood ratio contingency tests (G statistic) of biophysical factors and endemic plantsa (continued)

Biophysical factor Value of G df P

Rhyolite

Sand

Sedimentary or metamorphic

Slides and talus

Tuff

(Serpentine was omitted because

of small sample sizes) +

Landform: 7.111 3 0.05 < P < 0.10 +

Cliff

Ridgetop

Scablands

Valley bottom

Soil pH: 3.075 1 0.05 < P < 0.10 +

Acidic to neutral (combined

because of small sample sizes)

Alkaline

Soil texture: 1.363 2 0.50 < P < 0.75

Coarse (sand)

Medium

Fine

Soil depth: 3.896 3 0.25 < P < 0.50

Very shallow (<25 centimeters)

Shallow (25 to 50 centimeters)

Moderately deep (51 to 102

centimeters)

Deep to very deep (>102

centimeters)

Soil moisture:b 6.434 3 0.05 < P < 0.10 +

Andic

Udic

Ustic

Xeric

Soil temperature:b 0.144 2 0.90 < P < 0.95

Cryic (0 to 8 oC)

Frigid (diff. mean summer/

winter ≥ 5 °C)

Mesic (8 to 15 oC)

a The null hypothesis for each test was that biophysical characteristics of locally and regionally

endemic rare or potentially rare vascular plant species do not differ from those of nonendemic rare

or potentially rare species.

b See Harvey and others (1994:3) for definitions of terms.

30
Figure 17—Number of lichen, bryophyte, and vascular plant species groups by dispersal

mode.

Figure 18—Number of rare or potentially rare fungi

and vascular plant species by dispersal mode.

31
Figure 19—Number of rare or potentially rare vascular plant species by dispersal mode, showing (A)

dispersal by means of vertebrates and (B) dispersal by means of growth or reproduction.

32
including many more trees, shrubs, and other

forms—are likely to be very different.

Plant pollinators—Along with assuring suitable

habitat conditions and protection of specific locations, successful pollination of the rare or potentially rare vascular plants of the basin assessment

area may be one of the most important ecological

factors that can help ensure their long-term viability. Pollination ecology of these species, however, is essentially unstudied and unknown. What

follows are general patterns from initial presumptions as to pollination ecology. These patterns

need to be tested with field studies.

As Buchmann and Nabhan (1996) note, plantpollinator relations exemplify connections between endangered species and threatened habitats. Human-induced changes in pollinator populations—caused by overuse of chemical pesticides, unbridled development, and conversion of

natural areas into monocultural cropland—can

have a ripple effect on disparate species, ultimately leading to a “cascade of linked extinctions” (Buchmann and Nabhan 1996).

Pollinators also can be influenced by habitat fragmentation, sometimes differentially benefitting

exotic insect pollination vectors to the detriment

of native vectors. Aizen and Feinsinger (1994)

studied native and honey bee (Apis mellifera; an

exotic) pollinators of native flowering plants in

subtropical dry forest of northwestern Argentina.

They found that habitat fragmentation adversely

affected native flower visitors but facilitated the

use of floral resources by honey bees.

Loss of pollinators also can result in different selection pressures for reproductive strategies of

surviving plants. In a modeling study of an isolated population of Primula sieboldii, Washitani

(1996) reports that loss of pollinators may result

in strong fertility selection for a self-fertile,

homostyle morph. He concluded that active management of plants with absent or moribund pollinators could include hand pollinations and reintroduction and reestablishment of suitable pollinator populations. This would help maintain a

more natural gene pool in future plant

generations.

In the basin assessment area, by far most of the

rare or potentially rare vascular plants are pollinated by invertebrates (66 percent of plant species). Additional modes of pollination include

autogamous pollination (10 percent), wind (5 percent), hummingbirds (2 percent), and generalist

(1 percent), and others are unknown (16 percent)

(fig. 20). Most plants are serviced by multiple

pollinator species. For example, Eriogonum

prociduum (prostrate buckwheat) likely is pollinated by beetles (Cleridae: Coleoptera) but

also can be pollinated by flies (Tachinidae,

Sarcophagidae, Bombyliidae: Diptera), wasps (six

families: Hymenoptera), and moths (Sesiidae:

Lepidoptera).4

Of the plants pollinated by invertebrates, about

half are pollinated by bees, wasps, and allies

(Hymenoptera), with others pollinated by flies

(Diptera), butterflies and moths (Lepidoptera),

beetles (Coleoptera), spiders (Araneida), and

other insects (not specified) (fig. 21). Insects as a

whole play a vital function in reproduction and

viability of populations of angiosperms.

Long-term viability of many plant populations

likely is affected by viability of plant pollinator

and plant disperser mutualists. Many of these

relations are typically highly specific (Menges

1991). Terbourgh (1986) discussed keystone

mutualists, which are species that, if extinct, likely

would result in secondary extinctions of many

associated obligate species. Keystone mutualists

might exist among insect-plant pollinator relations in the basin assessment area (and possibly

also among plant-disperser relations as well).

Thus, understanding basic pollination and dispersal ecology of plants is key to projecting and

planning for long-term population viability.

Indicators—Plants and allies can serve as

bioindicators of environmental conditions and

ecosystem health. The concept is far from new,

beginning perhaps with Clemment’s (1920) usage

of plant indicators to denote vegetation communities.5 Others have extended the concept in various

directions. For example, Nygaard (1949) indexed

the trophic health and degree of eutrophication of

water bodies by calculating the ratio in number of

algae species of different taxonomic groups.

4 Source: OSDA-BLM Challenge Cost Share Project 89-6.

5 Actually, well into prehistory, many indigenous peoples

likely used plants to locate underground water, game

animals, and good places to build camps. Modern science

does not have the corner on observing nature.

33
Figure 20—Number of rare or potentially rare vascular plant species by type of pollinator.

Figure 21—Number (in parentheses) of rare or potentially rare vascular plant species by type of

invertebrate pollinator.

34
Westveld (1954) used plants to index site quality.

And, Anderson (1986) (and many others) identified plant indicators of range condition.

The Plant Task Group of the Science Integration

Team identified a number of lichen, bryophyte,

and vascular plant groups and vascular plant

species that can serve to index specific environmental conditions of the basin assessment area.

Such environmental conditions include air quality, carbonate levels, water flow levels (low,

high), high nitrogen conditions, metal-rich rock

conditions, old-growth forest conditions, soil

temperature (low, high), and soil texture (coarse,

fine). Specific indicators, and the environmental

condition for which they may be useful as an

indicator, are listed in appendix B.

Overall, many lichens can be used to index air

quality (fig. 22; see appendix B), a function

already finding utility in management of many

forests and other geographic areas (for example,

Geiser and others 1994, Stolte and others 1993).

Lichen groups along with bryophyte groups also

can be used to index most of the other categories

of environmental conditions listed above (fig. 22;

see appendix B). In addition, vascular plant

groups and rare or potentially rare vascular plant

species can be used as bioindicators of alkaline

(carbonate) soil conditions, overgrazing, metalrich rock conditions, old growth, low soil temperature, coarse soil textures, and other conditions (fig. 22; appendices B and C).

Nonvascular plant indicators of old-growth forest conditions may be of particular interest to

managers. Tibell (1992) identifies crustose

lichens as indicators of temporal continuity of

old boreal conifer forests. The term temporal

continuity refers to the degree to which specific

kinds of habitats—in this case, old boreal conifer

forests—continuously occupy a local geographic

area over time. Some studies outside the basin

assessment area have indicated that interrupting

the temporal continuity of old forests has led to

loss of sensitive species closely associated with

such habitats because there was insufficient time

or lack of sources for recolonization.

Lichens also are identified that indicate forest

stand temporal continuity in northern boreal forests (Esseen and others 1996, Nilsson and others

1995, Selva 1994). Pike and others (1975) identify

epiphytic lichens and bryophytes associated with

old-growth forests of western Oregon. Söderström

and Jonsson (1992) identify bryophytes that indicate fragmentation of old forests; that is, loss of

old-growth bryophyte species in the more isolated

forest fragments. In the current study, we identified 15 lichen groups and 9 bryophyte groups that

may indicate old-growth forest conditions in the

basin assessment area (see appendices B and C).

The rotten log and tree base lichen group in

particular indicates ecological continuity of old

growth.

Many of the bioindicator species and groups identified here, including those of old-growth forests,

can be useful to managers interested in quickly

surveying potential environmental conditions

without investing in long-term ecological research

studies. Such indicators can serve to track effects

of alternative management activities, such as variations in grazing, timber harvesting, or restoration regimes.

Invertebrates

Number of species—In the basin assessment

area, some 24,270 species of macroinvertebrates

are estimated to exist (Marcot and others 1997).

This figure extrapolates for species not yet discovered in the basin assessment area and some

not yet described, and comprises about 770

mollusks (425 freshwater gastropods, 35 freshwater bivalves, 30 slugs, and 280 land snails;

T. Frest in Marcot and others 1997) and 23,500

arthropods, including insects. At present, only

3,780 species of macroinvertebrates (16 percent

of all estimated species) have actually been reported from the basin assessment area, including

380 mollusks (200 freshwater gastropods, 30

freshwater bivalves, 25 slugs, and 125 land snails;

T. Frest in Marcot and others 1997) (49 percent

of all estimated mollusks) and 3,400 arthropods

(14 percent of all estimated arthropods). An additional, large set of micro-organisms, including

bacteria, protozoa, rotifers, viruses, and other

taxa, occur. They may number in the hundreds

of thousands of species but are mostly unknown

and are treated here only by taxonomic groups.

35
The SER database lists only a small example set

of 346 invertebrate species. Of these, environmental correlates and ecological functions are

denoted for 195 individual species of arthropods

and 11 species groups of micro-organisms (6 species groups of soil bacteria, 1 group of soil protozoa, 1 group of soil rotifers, and 3 groups of soil

nematodes). Thus, the analyses that follow provide only examples of invertebrate species and

micro-organism groups.

Dispersal modes and KEFs—Even for the relatively tiny set of example species explicitly

included in this assessment, various dispersal

modes and related KEFs were identified. Dispersal modes included various forms of independent

locomotion (crawling and other flightless terrestrial movement), sedentary habits, soil digging,

aerial movement (flying, ballooning, and drifting), budding, and phoresis (dispersal using other

host species).

Phoretic dispersal exemplifies some interesting

relations that have evolved among species. Phoretic dispersal has appeared in many microorganisms and invertebrates, including bacteria,

rotifers, protozoa, nematodes, and insects (table

7). Host species can include seeds, plants, organic

debris, other insects, birds, mammals, and even

humans. Soil bacteria are ubiquitous and disperse

by means of most of these hosts. Rotifers and soil

protozoa can hitch rides on arthropods, small

mammals, birds, and even soil nematodes.

Bacterial-feeding soil nematodes in turn can

travel by arthropods, birds, small mammals, and

even by humans via tools that redistribute soil,

such as shovels.

Figure 22—Number of lichen, bryophyte, and vascular plant species groups by bioindicator category. See appendices B and C for lists of groups, and text for explanation of categories.

36
Table 7—Examples of micro-organisms and invertebrates that disperse by phoresis (facultative associations among species, such as

being carried and dispersed by other organisms)Scientific name or Common name orgeneral species group specific species group Dispersal mode Dispersal requirementsBacteria:

Erwinia cartovora Bacterial pathogens Carried by seeds, plant debris, or alternative hosts, Presence of bacterium in diseased plant or animal

birds, or humans material; dispersal agent often required to move the

pathogen from diseased host to susceptible host

Microcoleus Beneficial bacteria Phoretic on arthropods or small mammals and birds Presence of the bacterial/lichens in undisturbed soil;

feeding by dispersal agent

Bacillus thermophilus Competitive bacteria Phoretic on arthropods or small mammals or birds, or Presence of bacterium in undisturbed soil; feeding by

by inoculation dispersal agent

Pseudomonas fluorescens Competitive bacteria in Phoretic on arthropods or small mammals or birds, or Presence of bacterium in undisturbed soil; feeding by

the rhizosphere by inoculation dispersal agent

Klebsiella planticola Bacteria: N-immobilizers Phoretic on arthropods or small mammals or birds, or Presence of bacterium in undisturbed soil; feeding by

by inoculation dispersal agent

Rhizobium melliotii Bacteria Phoretic on arthropods or small mammals or birds, or Presence of bacterium in undisturbed soil; feeding by

by inoculation dispersal agent

Rotifers:

Rotifers Rotifers: no info about Phoretic on arthropods, small mammals, birds, or Presence in undisturbed soil; presence of dispersal

rotifer spp. in basin nematodes agent

Protozoa:

Amoeba amoeba Protozoa: bacterial-predators Phoretic on arthropods, small mammals, birds, or Presence in undisturbed soil; presence of dispersal

nematodes agent

Nematodes:

Acrobeloides Bacterial-feeding nematodes Phoretic on arthropods, small mammals, or birds Presence in undisturbed soil; presence of dispersal

agent

Aphelenchus avenae Fungal-feeding nematodes Phoretic on arthropods, small mammals, or birds Presence in undisturbed soil; presence of dispersal

agent

Pratylenchus penetrans Plant-feeding nematodes Phoretic on arthropods, small mammals, birds, or Presence of undisturbed soil; presence of dispersal

human tools (shovel) agent

37
Table 7—Examples of micro-organisms and invertebrates that disperse by phoresis (facultative associations among species, such as

being carried and dispersed by other organisms) (continued)Scientific name or Common name orgeneral species group specific species group Dispersal mode Dispersal requirementsFungi-related:

Hysterangium Ectomycorrhizal mat- Carried by nematodes, arthropods, or small mammals Dispersal agents

forming fungi

Rhizoctonia solani Fungal pathogens Carried by seeds, plant debris, or alternative hosts, Presence of bacterium in undisturbed soil; feeding by

birds, or humans dispersal agent

Glomus mossae Vesicular fungi Carried by seeds, plant debris, arthropods, ants, or Dispersal agents

wind

Penicillium citrinum Saprophytic fungi Phoretic on arthropods, small mammals, birds, or by Presence of bacterium in undisturbed soil; feeding by

inoculation dispersal agent

Insecta:

Cimex latipennis Bed bug (Cimicidae: Flightless—disperse only on host animal Presence of bats when proper life stages of bug

Hemiptera) available

38
Phoretic invertebrates play various roles in contributing to trophic structures and ecological

processes of ecosystems. Many phoretic invertebrates consume soil micro-organisms and are

prey for other consumers; they aid soil nitrogen

cycling through nitrogen immobilization and by

acting as a source for nitrogen mineralization,

which can be vital functions in forest nitrogen

conservation (Davidson and others 1992, Miller

and others 1989). A few phoretic invertebrates

are parasites or pathogens, or cause disease. Phoretic invertebrates also include soil-oriented species whose functions aid soil structure, enhance

soil stabilization, detoxify xenobiotics, sequester

(accumulate) soil metals, and influence vegetation succession and presence and diversity of

plant species.

The SER database afforded the opportunity to explore some examples of how dispersal modes of

invertebrates relate to their KEFs. Invertebrates

with independent locomotion provide many ecological services in their ecosystems. Most of these

example species included in the SER database

were insectivorous and provide food for other

insectivores, with a few being carrion feeders,

detrivorous, or herbivorous on trees or shrubs, or

feeding in water on algae or plankton. The four

sedentary species were folivores.

Species dispersing by digging enhance the physical structure of soil and turnover of soil nutrients

and layers. Earthworms are a prime example (see

discussion in Marcot and others 1997). In one

study of eight species of earthworms in a Kansas

tallgrass prairie, James (1991) reported that total

annual soil consumption by all earthworms was 4

to 10 percent of organic matter in the top 15 cm;

100 to 300 percent of plant annual belowground

production passed through the earthworms each

year; and mineral nitrogen processed was 10 to

12 percent of annual plant nitrogen uptake, comparable to half of the input from precipitation,

whereas the phosphorus processed was equivalent to 50 percent of annual uptake. Native and

introduced earthworm species, however, do not

necessarily perform the same functions. James

found that exotic, introduced earthworm species

have a negative effect on soil turnover and nutrient mineralization because of the lower soil

throughout and their relative intolerance of

summer soil temperatures, as compared with

the native earthworms.

Invertebrate species that disperse by flying, ballooning, or drifting cover an even wider array of

trophic functions than those with independent

locomotion, and include folivores, spermivores,

sap feeders, root feeders, insectivores, soil microorganism feeders, carrion feeders, aquatic herbivores, detritovores, and moss feeders. Many of

these aerial-dispersing species (many Hymenoptera) serve as important pollination vectors critical

for reproduction of flowering plants. They also

include forest beetles, some of which can change

the structure of terrestrial and aquatic habitats by

causing tree mortality, breaking down standing

dead trees and large down logs, creating canopy

gaps in forest stands, and causing increased

streamflow. Thus, providing suitable conditions

and means of dispersal for invertebrates also provides for many ecological processes and species

functions in the ecosystem.

Disease and parasites—The ecology of disease

and parasites is not well studied in the basin assessment area and is poorly represented in the

SER database. Disease can strongly mediate biodiversity and viability of many kinds of wild

plants and animals and needs to be considered in

wildlife conservation biology programs (Dobson

and Mat 1986, May 1988, Scott 1988). Aguirre

and Starkey (1994) conclude that proximity of

domestic ungulates to wild ungulates can transmit

disease and parasites and complicate wild ungulate management; they cite several examples,

including bovine brucellosis (Brucella abortus) in

bison (Bison bison) and Rocky Mountain elk in

Yellowstone National Park, and lungworm

(Pasteurella spp.)-pneumonia complex in bighorn

sheep (Ovis canadensis) in several Western national parks (also see Peterson 1991). Simonetti

(1995) cites other similar examples, and

Cunningham (1996) warns of secondary transmission of parasite pathogens to wildlife during translocation projects. Disease transmission can exacerbate recovery of endangered species (Thorne

and Williams 1988) and is a controversial facet of

wild game ranching. Certainly, too, human health

concerns warrant attention to wild pathogens,

such as transmission of lyme disease by ticks

(Ixodes scapularis) throughout the basin assessment area (Ostfeld and others 1995).

Parasites, however, also play ecological roles in

ecosystems and serve as an ecological “glue” to

bind many species’ relations. Windsor (1995,

39
1996) provides many fascinating examples of

this, including the following. Hybrid cottonwoods

(Populus fremontii x P. angustifolia) are more

susceptible to aphid parasites than are their parent

species. Thus, aphids concentrate in the hybrid

zone between the parents. This aphid sink discourages aphids from adapting to the more numerous parents, and it keeps the parents as separate species. Peterson (1991) calls for controlled

experiments, or at least an adaptive management

approach (learning by trial), to provide a sounder

scientific basis for management of parasite-host

interactions.

Many invertebrate parasite-host relations await

discovery. For example, Boonstra and others

(1980) found that botfly parasitism can affect

vole populations. Dobson (1988) suggests that

parasites can be used as an effective conservation

biology weapon to control introduced mammals

threatening fragile isolated or island biota.

Vertebrates

Number of species—In the basin assessment

area, 468 species of vertebrates (excluding fish)

are both estimated and known to exist (Marcot

and others 1997). These include 26 species of

amphibians, 27 reptiles, 283 birds, and 132 mammals. An additional 79 bird species are accidental

or casual species in the basin assessment area.

The SER database lists 476 species of vertebrates,

consisting of all the known species and including

291 species of birds (excluding the accidentals or

casuals, but including 8 bird species that likely

occur only sporadically in the basin assessment

area). Thus, the analyses that follow provide a full

accounting of all regularly occurring vertebrate

species.

Endemic species—Findings on vertebrate species

endemism within the basin assessment area are

presented in Marcot and others (1997). To recap,

two amphibian species are local endemics: Idaho

giant salamander (Dicamptodon aterrimus) and

Larch Mountain salamander (Plethodon larselli);

and seven other amphibian species are regional

endemics: northwestern salamander (Ambystoma

gracile), Cope’s giant salamander (Dicamptodon

copei), Dunn’s salamander (Plethodon dunni),

Coeur d’Alene salamander (P. idahoensis), Cascade torrent salamander (Rhyacotriton cascadae),

red-legged frog (Rana a. aurora), and Cascades

frog (R. cascadae). The locally endemic amphibian species occur in 9 vegetation cover types,

and the regionally endemic species occur in 20.

Locally or regionally endemic amphibians are

particularly rich (more than six species) in several

montane forest vegetation cover types, including

Pacific silver fir/mountain hemlock forest, mountain hemlock forest, interior Douglas-fir forest,

and western redcedar/western hemlock forest.

Other vegetation types, however, also may be

necessary to ensure the long-term persistence

and distribution of some of these species.

No reptile was determined to be locally endemic

or regionally endemic. Many reptile species occur

only along the margins of the basin assessment

area and beyond, such as in the Great Basin and

Klamath Basin to the south.

There are no locally endemic birds, and four are

regionally endemic: sage grouse (Centrocercus

urophasianus), rufous hummingbird (Selasphorus

rufus), northern spotted owl (Strix occidentalis

caurina), and Columbian sharp-tailed grouse

(Tympanuchus phasianellus columbianus).

Collectively, these taxa occur widely throughout most vegetation cover types, although specific ranges of some of these taxa (especially

Columbian sharp-tailed grouse) are quite limited

within the basin assessment area.

Among mammals, local endemics include two

species (Idaho ground squirrel (Spermophilus

brunneus) and Washington ground squirrel

(S. washingtoni)) and two subspecies (potholes

meadow vole (Microtus pennsylvanicus kincaidi)

and White Salmon pocket gopher (Thomomys

talpoides limosus)). There may be other locally

endemic subspecies of mammals not recognized

in this study. Some 22 other mammal species are

regional endemics (see Marcot and others 1997

for list). Collectively, local endemics and regional

endemics occur widely throughout most vegetation cover types of the basin assessment area but

are concentrated somewhat more in drier forest,

shrub, and herb communities. Vegetation cover

types that are particularly rich in locally or

regionally endemic mammals (>12 species) include mixed-conifer woodland, big sagebrush

(Artemisia vaseyana), native forb, mountain big

sagebrush, and low sage. High numbers of

endemics occur in mixed-conifer forests probably

40
because such forests occupy a wide range of

moisture and temperature conditions and their

overall species diversity is typically high (Harvey

and others 1989, Sayler and Martin 1996).

The number of species that are local and regional

endemics among amphibians, reptiles, birds, and

mammals was not significantly different than the

number of all nonendemic vertebrates by taxonomic class. That is, local and regional endemics

were not more apt to be of any particular vertebrate class (Marcot and others 1997).

Species by percentage of range within the

basin assessment area—A complementary way

of considering patterns of endemism is the percentage of a species’ range that lies beyond the

basin assessment area. We identified five categories for percentage of range: (A) nearly all

(>79 percent) of the species’ range lies beyond the

basin assessment area; (B) more than half but

not nearly all (70 to 79 percent) lies beyond; (C)

about half (69 to 40 percent) lies beyond; (D) less

than half (20 to 39 percent) lies beyond; and (E)

only a very small portion (<20 percent) lies

beyond.

By taxonomic class, about two-thirds of the amphibian species have more than half their ranges

beyond the basin assessment area (range categories A and B); only two amphibian species (the

two local endemics) are locally confined (range

category E) (fig. 23). Reptiles and birds are even

more broadly distributed. Among mammals,

about 95 percent of the species have half or more

of their range beyond the basin assessment area

(range categories A, B, and C); and only six taxa

Figure 23—Number of vertebrate species by taxonomic class, by proportion of species’ range lying

outside the basin assessment area. Species range categories: A = nearly all (80 to 99 percent) of

the overall species range occurs outside the assessment area; B = more than half, but not nearly

all (60 to 79 percent) occurs outside; C = about half (40 to 59 percent) lies outside; D = less than

half, but not only a very small portion (20 to 39 percent) lies outside; and E = only a very small

portion (0 to 19 percent) lies outside.

41
have their range mostly or entirely within the

basin assessment area (range category E). These

six mammal taxa include all four local endemics

plus two other sciurids: Uinta ground squirrel

(Spermophilus armatus) and red-tailed chipmunk

(Tamias ruficaudus).

Species that migrate and spend part of their

year elsewhere but that restrict their distribution during one part of the year to a specific region (such as the basin assessment area), are

called semiendemic species by Garza (1996).

Semiendemics can include Neotropical migratory

birds and other species that occur only seasonally

in a given region. Garza argues that species that

are confined to a specific geographic region, even

if only for a portion of the year, still should be

considered as restricted to that area for purposes

of habitat conservation. Because our range maps

show neither season of use nor migration range

beyond the basin assessment area, our designation of migratory species into endemism and percentage of range classes approximates the concept

of semiendemics. Thus, some migratory species

might occur as semiendemics in the basin assessment area but rank only as range categories A, B,

or C.

Another way of looking at abundance patterns

among range classes is shown in figure 24. Most

species that extend beyond the basin assessment

area (range categories A, B, and C) are birds and

mammals, with a smaller proportion of amphibians and reptiles. Those somewhat more confined

to the basin assessment area (range category D)

are all mammals, and those most confined to the

basin assessment area (range category E) are amphibians and small mammals.

We were surprised to find so few vertebrates, especially species with small body size, to be local

or regional endemics, and so few vertebrates with

ranges mostly confined to the basin assessment

area. It is evident that the administrative and hydrologic boundaries of the basin assessment area

do not particularly circumscribe unique vertebrate

communities and species ranges. This is because

the basin assessment area is “land-locked,” and

much of its ecological character along the margin

spills into adjacent areas, including the western

slope of the Cascade Range in western Washington and Oregon, the Klamath Basin and Great

Basin in northern California and Nevada, the

Greater Yellowstone Ecosystem in Wyoming, the

northern glaciated mountains in the Canadian

Rocky Mountains, and the Okanogan Highlands

in the Okanagan6 Desert of British Columbia,

Canada. Marcot and others (1997) list vertebrate species closely associated with many ERUs,

and many of these assemblages consist of species

that range well beyond the basin assessment area

in many of these directions.

Abundance, breeding status, and migration

status of birds—Abundance, breeding status,

and migration status of birds in particular were

categorized by state and for the entire basin assessment area. The total number of birds (including accidentals and casuals) occurring by

state was remarkably constant, ranging between

only 360 and 362 species. The total number of

breeding birds (migrants plus residents) by state

averaged 221 species and ranged from 192 species in Wyoming to 242 species in Oregon. (It

should be remembered that these species tallies

refer only to those portions of these states that lie

within the basin assessment area.) This breedingbird richness is quite close to that suggested by

Huston’s (1994:25) plot of latitudinal gradient

in species richness of breeding birds in North

and Central America. The basin assessment area

lies between latitude 42o and 49o N., for which

Huston’s bird species richness curve predicts

about 200 species (but the data scatter on the

curve suggests upward of 300 species

empirically).

For the basin assessment area as a whole, most

bird species were ranked as common, with fewer

species uncommon or rare, fewer still as abundant, and the fewest as irregular (fig. 25).7 This

general bird abundance pattern also held for most

of the five western states included in this part of

the assessment (Washington, Oregon, Idaho,

6 A word on spelling: the United States has the Okanogan

Highlands and Canada has the Okanagan Desert, but they do

constitute a single shared ecosystem.

7 If the abundance categories can be taken as approximate

octaves or geometric series, then the bird species

abundance histograms match well those reported in other

north temperate ecosystems, such as nesting bird species in

Quaker Run Valley, New York (Krebs 1978:452). Such

species abundance patterns are somewhat stable attributes

of continental communities.

42
Figure 24—Percentage of all vertebrate species by taxonomic class, showing proportion

of species’ range lying outside the basin assessment area (see fig. 23 for description of

species range categories).

Figure 25—Number of bird species by abundance category (see text for definitions of

categories).

43
Montana, and Wyoming; data for Nevada and

Utah were not available for those small portions

present in the basin assessment area), with a few

deviations (fig. 26). Most of the irregular species

occur in western Montana, which is located

toward midcontinent and which picks up many

species found more commonly in other, adjacent

continental ecosystems. Examples of irregular

species in western Montana are McCown’s longspur (Calcarius mccownii), which is more of a

prairie species farther east, black-throated blue

warbler (Dendroica caerulescens), which occurs

more commonly in Eastern U.S. states, and hoary

redpoll (Carduelis hornemanni), which is an irregularly irruptive species from northern latitude

boreal and subboreal forests. On the other hand,

likely for the same reasons of being more central

in the continent, western Montana has the greatest

number of abundant species, and northwestern

Wyoming has the greatest number of common

species in the basin assessment area (fig. 26).

As for breeding and migration status of birds

throughout the basin assessment area, most species are resident (36 percent) or summer breeders

(26 percent), with fewer species occurring as migrants (14 percent), winter only (7 percent), or

nonbreeding summer residents (2 percent) (the remaining 15 percent were not specified) (fig. 27).

Proportions of bird species in these breeding and

migration classes generally were consistent across

the states but with a decreasing proportion of

residents and increasing proportion of migrants

toward midcontinent (fig. 28).

Tallying number of bird species by abundance

status and by breeding and migration status

revealed interesting geographic trends by state

(fig. 29). Throughout the five states included in

this part of the analysis, most of the abundant,

common, and uncommon bird species were residents and summer breeders. Far fewer abundant

bird species occurred in Idaho and Wyoming than

in the other three states. Rare bird species included various mixes of breeding status categories

by state with no clear geographic pattern,

Text continues on page 48

Figure 26—Number of bird species by abundance category and U.S. state, within the

basin assessment area (see text for definitions of categories).

44
Figure 27—Number of bird species by breeding status category, within the basin

assessment area (see text for definitions of categories).

Figure 28—Percentage of bird species listed by breeding status category and U.S. state,

within the basin assessment area (see text for definitions of categories).

45
Figure 29—Number of bird species listed by abundance category, breeding

status category, and U.S. state, within the basin assessment area (see text

for definitions of categories).

Figure 29—continued

46
Figure 29—continued

Figure 29—continued

47
although Wyoming and Washington had more

rare migrants than did other states. Among all

five states, irregular species consistently comprised mostly migrants. Because of their geographic mobility, migrants, especially longdistance migrants, are more apt to appear irregularly in individual states than are birds of

other breeding status categories.

Brood parasitism—Among vertebrate parasites,

brown-headed cowbirds (Molothrus ater) are nest

parasites of Neotropical migratory songbirds and

are a factor in their decline (Brittingham and

Tempe 1983, Rothstein and others 1980; see

Marcot and others 1997 for a list of host species of the basin assessment area). Not all hosts

may respond the same, however. Uyehera and

Narins (1995) experimentally demonstrate that

willow flycatchers (Empidonax traillii) can differentially recognize female and male cowbirds

near their nests, and that the behavior of the flycatchers acts as a counteradaptation to thwart

brood parasitism.

Species function profiles—Numbers of species

with selected KEFs were tallied by terrestrial

vegetation community. We term these depictions

“species function profiles” (fig. 30), which depict

variations in functional redundancy among terrestrial vegetation communities (table 2). For this

analysis, we selected a range of KEFs that seemed

particularly pertinent to vertebrates and for which

variations in numbers of vertebrate species among

communities might prove instructive (table 8).

We explored patterns of two trophic relations

functions. The first was presence of heterotrophs

(table 8), which can include primary consumers

or herbivores, secondary consumers or primary

predators, tertiary consumers or secondary predators, omnivores, carrion feeders, cannibals, and

coprovores (feed on fecal material). As expected,

the number of vertebrate heterotroph species

did not differ significantly among terrestrial vegetation communities (fig. 31a) because heterotrophy is a generalized function of all vertebrate

species (380).

We then narrowed the focus to carrion feeders,

which include at least seven species: Great Basin

spadefoot (Spea intermontana), which also feeds

on insects, small invertebrates, and sometimes

each other (cannibalism); coyote (Canis latrans),

an opportunist scavenger for which carrion is

most important during winter; wolverine (Gulo

gulo); turkey vulture (Cathartes aura), whose

feeding behavior significantly contributes to decomposition of carcasses; bald eagle (Haliaeetus

Figure 29—continued

Text continues on page 52
Figure 30—The general form of species function profiles showing functional redundancy and total species functional diversity

in a community. Each of the KEFs in a community has a number of species with that function. Species function profiles are

graphs displaying the numbers of species with each specific

KEF across communities. Functional redundancy is the number

of species with a specific KEF in a given community. Total species functional diversity is the number of KEFs times the mean

number of species per KEF (adapted from Huston 1994:5).

Table 8—Codes used in the species function profiles (fig. 31) for key ecological functions (KEFs)

of speciesa

KEF code Description of KEF

1.2 Heterotrophic consumer (380)

1.2.5 Carrion feeder (7)

2.1 Ungulate herbivory (may influence rate or trajectory of vegetation succession and

presence of plant species) (15)

3.1 Nutrient cycling relations: aids in physical transfer of substances for nutrient

cycling (C,N,P, other) (32)

4.1 Interspecies relations: potential insect population control (22)

4.2 Interspecies relations: vertebrate population control (10)

4.3 Interspecies relations: pollination vector (6)

4.4 Interspecies relations: transportation of seed, spores, plant or animal disseminules (153)

4.8 Interspecies relations: primary cavity excavator in snags or live trees (17)

4.9 Interspecies relations: primary burrow excavator (fossorial) (39)

6.1 Soil relations: physically affects (improves) soil structure, aeration (typically by

digging) (10; combined with KEF 6.2)

6.2 Soil relations: aids general turnover of soil nutrients and layers (10; combined with KEF 6.1)

7 Wood relations (breaks down wood) (4)

a Total number of vertebrate species with each KEF within the basin assessment area is shown in parentheses. Taken from a

longer classification presented in Marcot and others (1997).

49
Figure 31—Vertebrate species function profiles. These show the

number of vertebrate species that have specific KEFs, listed by terrestrial vegetation community, within the basin assessment area. See

table 8 for description of KEFs. (A) KEF 1.2 = heterotrophic consumer,

KEF 4.4 = transportation of plant or animal disseminules; (B) KEF

1.2.5 = carrion feeder, KEF 2.1 = herbivory; (C) KEF 3.1 = nutrient

cycling, KEF 4.1 = potential insect control, KEF 4.2 = vertebrate

population control; (D) KEF 4.3 = pollination vector, KEF 4.8 = primary

cavity excavator; and (E) KEF 4.9 = primary burrow excavator, KEF 6.1

or 6.2 = soil structure relations, KEF 7 = wood relations (breaks down

wood). Note that three terrestrial vegetation codes included in table 2

are not displayed here; these are urban, water, and rock-barren

communities.

Figure 31—continued

50
Figure 31—continued

Figure 31—continued

51
leucocephalus), which can also take live prey;

common raven (Corvus corax); and gray jay

(Perisoreus canadensis), an omnivore. Other vertebrate species doubtless feed on carrion as well.

Presence of the seven species listed here was

more variable among terrestrial vegetation communities (fig. 31b) than was the general function

of heterotrophy. The fewest number of carrionfeeders—that is, the least functional redundancy

of the vertebrate community for this particular

ecological function—occurred in early seral ponderosa pine forest and early seral subalpine forest

vegetation communities, and reached its maximum in late seral forests and woodlands.

Herbivory is another subset of heterotrophy, and

an ecological function that displayed a profile different from that of carrion feeding. Herbivory

reached its greatest functional redundancy in early

seral montane forest and in upland shrub vegetation communities (fig. 31b). As an ecological

force, herbivory can alter the vegetation structure

and succession of habitats. For example, moose

(Alces alces) heavily browse early seral species

such as aspen and willow, which can result in increases of conifers in habitats otherwise dominated by deciduous shrubs and trees (also see

Pastor and others 1993). In semipalustrine habitats, plant communities can be altered by herbivory of several species of waterfowl, including

greater white-fronted goose (Anser albifrons),

brant (Branta bernicla), snow goose (Chen

caerulescens), and Ross’ goose (C. rossii).

Montane voles (Microtus montanus) and meadow voles (M. pennsylvanicus) can modify grassland structures through herbivory. Mule deer

(Odocoileus hemionus), white-tailed deer (O.

virginianus), Rocky Mountain elk (Cervus

elaphus nelsonii), and mountain goat (Oreamnos

americanus) can have major influences on composition and succession of vegetation through

herbivory. Mountain goats can alter alpine and

some subalpine plant communities. Herbivores

can also interact with pollinators and affect plant

reproduction and the evolution of plant traits

(Brody 1996).

Another KEF is that of nutrient cycling relations,

particularly species that substantially aid in physical transfer of substances for nutrient cycling.

Essentially, all organisms cycle nutrients. At least

some 32 vertebrate species, however, affect nutrient cycling substantially in their environments

(see Marcot and others 1997 for discussion). The

vertebrate species function profile for generalized

nutrient cycling relations suggests a fair amount

of variability of functional redundancy among

communities (fig. 31c). Communities with the

greatest redundancy in nutrient cycling functions among vertebrate species include late seral

stages of most forest types, upland and riparian

woodlands, and early seral montane forest. ComFigure 31—continued

52
munities with the least redundancy include agricultural, alpine, exotic, upland herb and shrub,

and riparian herb and shrub communities, possibly because of overall lower vertebrate species

richness in at least some of these communities.

We developed species function profiles for six

categories of interspecies relations: potential insect population control, vertebrate population

control, pollination vectors, transportation of

plant or animal disseminules, primary cavity excavation, and primary burrow excavation (fig.

31). Some 22 species of insectivorous amphibians, birds, and mammals have the potential for

controlling some nonirruptive insect populations;

some woodpeckers can dampen the amplitude of

irruptive insect populations (Koplin 1969). Functional redundancy of potential insect control was

greatest in upland and riparian woodlands and

least in upland and riparian herbland, exotic, and

agricultural communities.

Predatory activities of at least 10 species can control vertebrate populations. The species include

five mammalian carnivore predators and two raptors. Examples include American kestrel (Falco

sparverius), which may aid in population control

of some insects and rodents, and American

badger (Taxidea taxus), which may help control ground squirrel populations. Redundancy

of vertebrate population control was less variable

among communities but reached its peak in upland and riparian shrub, upland woodland, early

seral montane forest, and—interestingly—

alpine communities; and was least in most mid

and late seral forest communities.

At least six birds—five hummingbirds and an

oriole—serve as pollination vectors for flowering

plants (see Marcot and others 1997 for discussion). This species assemblage did not show

much variation in redundancy among vegetation

communities (fig. 31d). Another, much larger set

of 153 species serve to transport plant or animal

disseminules; this set showed a fair amount of

variation in redundancy among communities,

with the greatest redundancy (most species) in

early seral montane forest, upland shrub, riparian

woodland, and upland woodland communities,

and the least redundancy in early seral ponderosa

pine forest and early seral subalpine forest (fig.

31a).

Primary cavity excavators—mostly woodpeckers,

nuthatches, and chickadees—number 17 species

(see Marcot and others 1997 for discussion) and

showed by far the greatest variation in functional

redundancy among vegetation communities of all

ecological functions explored here (fig. 31d). Primary cavity excavation reached its greatest redundancy by far in all late seral forests and upland

and riparian woodland communities; and its

lowest redundancy in agricultural, alpine, exotics,

upland and riparian herbland, upland and riparian

shrubland, and early seral forest communities.

Primary burrow excavators, a set of 39 species

(see Marcot and others 1997 for discussion), also

showed significant variation among communities.

The greatest functional redundancy was reached

in early seral montane forest, upland and riparian

shrub, and alpine communities, and the least functional redundancy was reached in late seral forest

communities, a rather inverse pattern to that of

primary cavity excavators (fig. 31e). As with

invertebrate soil excavators, among vertebrates

the primary burrow excavators can be important

to soil turnover and mixing of organic matter.

Gophers, for example, can process an enormous

quantity of soil material. In one study in coastal

San Diego County, California, Cox (1990) found

that the total soil mined by pocket gophers

(Thomomys bottae) in 10 Mima-type mounds

amounted to 8.23 Mg-1•ha-1

•yr

-1 and subsurface

deposition was 20.31 Mg-1•ha-1

•yr

-1, so that total

soil mining equalled 28.54 Mg-1•ha-1

•yr

-1.

Finally, soil relations and wood relations showed

relatively little variation in functional redundancy

among communities (fig. 31e). Soil relations

functions showed the greatest redundancy in early

seral montane forest, and upland and riparian

shrub communities, and the least redundancy in

early seral ponderosa pine forest and early seral

subalpine forest communities.

What general trends can be summarized from the

patterns of these selected ecological functions?

First, no one vegetation community harbors the

greatest redundancy of all ecological functions.

Most vegetation communities contribute to

greatest redundancy of some specific ecological

function. Thus, collectively, all vegetation communities combined help provide for redundancy

of all ecological functions. Exceptions seem to be

53
agricultural and exotic (and urban) communities,

which typically provide poorly for most functions explored here.

As a corollary, late seral forest communities seem

to provide the greatest redundancy in carrion

feeding, general nutrient cycling, and primary

cavity excavation functions. Late seral forest

communities provide poorly for vertebrate population control and primary burrow excavation

functions, which reach their peaks of redundancy

in other communities, principally early seral forest communities.

Alpine communities provide for high redundancy

in vertebrate population control and in primary

burrow excavation functions. Out of six mammal

species that regularly function as secondary burrow users (that is, use burrows created by primary

burrow excavators), however, only two species

(ermine [Mustela erminea] and long-tailed weasel

[M. frenata]) occur in alpine communities, and

these species occur widely throughout other vegetation communities as well. Thus, although alpine

communities provide for high functional redundancy of primary burrow excavation, it does not

provide for high redundancy of secondary burrow

use.

Upland and riparian herblands do not provide for

maximum redundancy of any of the functions explored here. Upland and riparian shrublands provide for high levels of redundancy of herbivory,

vertebrate population control, pollination, primary

burrow excavation, and soil-relations functions.

Upland and riparian woodlands provide for high

levels of redundancy of carrion feeding, general

nutrient cycling, potential insect population control, vertebrate population control, pollination,

and primary cavity-excavation functions. Thus,

shrublands and woodlands provide for complementary sets of ecological functions, just as do

late and early seral forest communities.

Early seral montane forest seems to provide for

an anomalously wide array of conditions and ecological functions of all forest types in the basin

assessment area. It provides for high redundancy

of general nutrient cycling, vertebrate population

control, pollination, primary burrow excavation,

and soil-relations functions, whereas early seral

ponderosa pine forest and early seral subalpine

forest communities do not. This may have to do

with the overall greater habitat and biotic diversity found in early seral montane forests than in

early seral stages of other forest types, but this

needs verification.

Which KEFs considered here are the most variable among terrestrial vegetation communities in

terms of number of vertebrate species (redundancy of species)? The answer is found by plotting

the standard error (SE) of number of species

among vegetation communities for each KEF. Results (fig. 32) show that, of the functions considered here, primary cavity excavation is by far the

most variable in functional redundancy of species

among all terrestrial vegetation communities. This

means that only specific vegetation communities

are likely to support most of the species with this

function. The next two functions with high variation in redundancy are the carrion feeder function

and the primary burrow excavator function. Thus,

the vegetation communities in which these mostvariable functions occur are likely to be important

for maintaining these species’ ecological functions within the basin assessment area. On the

other hand, KEFs with the lowest SEs include

transport of plant or animal disseminules, heterotrophic consumer, and pollination vector functions. These are the least variable in redundancy

and the most reliable among communities; none

of the vegetation communities is particularly critical for maintenance of these functions within the

basin assessment area (although specific composition of species with these functions may differ among communities).

Which terrestrial vegetation communities are the

most variable in terms of redundancy among ecological functions? The answer is found by plotting the SE of species richness (number of species) among KEFs for each vegetation community. Results (fig. 33) suggest that early seral

montane forest has the greatest variation, and

three communities (mid seral montane forest,

mid seral ponderosa pine forest, and mid seral

subalpine forest) have the least variation in redundancy among the ecological functions discussed above. This does not mean that lowvariation vegetation communities contain all

KEFs, but only that the functions they do provide

for tend to vary the least in number of species per

function; there is a greater “evenness” (relative

proportion) but not necessarily “richness” (number of different kinds) of species’ ecological

functions.

54
Overall, it is unclear how the degree of functional redundancy of KEFs, either among species

or among vegetation communities, specifically

affects long-term resiliency of ecosystems to perturbations of the environment and to perturbations of species community structures. A working

hypothesis might suggest that a greater redundancy promotes greater resiliency in ecosystem

ecological processes to stochastic environmental

events and to short-term or localized losses of

some species. This, however, is largely unstudied

in the basin assessment area. Our approach provides an analytic framework and a repeatable

means of posing such testable hypotheses on

ecosystem processes and functional redundancy

among communities.

Total species functional diversity—Huston

(1994) characterized total species functional diversity as the number of different ecological functions performed by all species in a community

times the mean number of species per function.

Figure 34 presents estimates of total species functional diversity of vertebrates for terrestrial vegetation communities of the basin assessment area

by using the ecological functions discussed above.

Over all communities, the greatest vertebrate

functional diversity is found in early seral montane forest, followed by upland woodlands and

riparian woodlands, and upland shrublands. Historically, most vegetation cover types associated

with early seral montane forest and upland woodlands have increased in total area in the inland

West since early historic times, whereas those of

riparian woodlands and upland shrublands have

decreased. Vertebrate species within these cover

types have shifted in relative abundance even if

vertebrate functional diversity has remained more

constant.

The lowest vertebrate functional diversity is

found in early seral ponderosa pine forest, early

seral subalpine forest, and agricultural lands

(fig. 34). The vegetation cover types associated

with early seral ponderosa pine forest communities have decreased in area since early historic

Figure 32—Variation in functional redundancy of KEFs among terrestrial vegetation communities,

displaying SE of number of species among vegetation communities, by KEF. Greater SE values

denote greater variation in functional redundancy among communities (see fig. 31 and table 8 for

descriptions of KEF codes).

55
times; those associated with early seral subalpine

forest communities have both increased and decreased; and agricultural lands have greatly increased. Interestingly, exotic vegetation communities do not rank particularly low in overall

vertebrate functional diversity (fig. 34), although

there are many vertebrate species that do not find

suitable conditions in such habitats (Marcot and

others 1997). Thus, levels of vertebrate functional diversity among vegetation comunities are

complementary to—not coincident with—patterns

of vertebrate species diversity (see Marcot and

others 1997 for species diversity descriptions).

To what extent can we expect functional redundancy among vertebrate species with the same

KEF? This question was addressed by Marcot

and others (1997) who conclude that general

functions, such as herbivory, cavity excavation,

and pollination, might be considered redundant if

performed by different species, but because each

species has its unique combination of habitat associations and life history patterns, no two species can be expected to be exactly interchangeable. Thus, patterns of species functional diversity presented here must be interpreted only as

broad-scale geographic and macroecological

trends and not interpreted as evidence for allowing any particular species loss or replacement in

communities.

Climate Change and Species at Risk

Results of climate change modeling suggest potentially complex local and regional climate responses in the basin assessment area (see Hann

and others 1997).8 Potential changes in climate

patterns among geographic locations within the

basin assessment area are not consistent in indicating warming and drying trends, as suggested

by Covington and others (1994) for the inland

Figure 33—Variation in functional redundancy of KEFs among functions, displaying SE of number

of species among KEF, by terrestrial vegetation community. Greater SE values denote greater

variation in functional redundancy among KEFs (see table 2 for descriptions of terrestrial

vegetation communities).

8 Output from only one climate model—RegCM2, a

regional climate model linked with a general circulation

model to test regional responses to 2xCO2—is currently

available for the basin assessment area. Hann and others

(1997; also see footnote 2 in chapter 3) summarized results

of that model along with knowledge of patterns that control

regional climate.

56
West,9 and by general circulation models for boreal forests of high latitudes. The growing season

could become +1 o

C warmer with slightly greater

spring precipitation (+5 percent) and slightly less

autumn precipitation (-5 percent). Above 1000 m

elevation, however, winter might experience

slight cooling (-1 o

C minimums and -2.5 oC maximums). This, along with significant increases in

winter precipitation (+30 to 50 percent) could

result in increased snow depth at high elevations,

although the general circulation models reported

by Covington and others (1994) suggest overall

regional warming trends. Below 1000 m elevation, winter might experience significant warming

(up to +3 o

C minimums and maximums) (see

footnote 2 in chapter 3).

Over all elevations, temporal changes in temperature and precipitation would not occur smoothly.

Thus, individual plant species adversely affected

would be those unable to adapt to changes or increases in interseasonal and interannual variation

in temperature and precipitation. Mostly, climate

changes and species-specific vegetation response

in the basin assessment area are poorly known

and not well modeled.

Given this uncertainty, we inspected the potential

response of species to one possible aspect of

climate change, namely, high-elevation warming

during the growing season. Knoll (1984) noted

that the effect of climatic change on extinctions

of plant species depends less on absolute changes

of mean annual temperature and precipitation, and

more on successful migration of populations over

generations to suitable areas. If suitable areas

do not exist, such as caused by warming trends

pushing upper elevation climate and life zones

into more restricted distributions (that is, into

areas of barren rock or ice, or even off the tops

of mountains), or if access to suitable areas is

blocked by dispersal barriers, then local extinction rates greatly increase. We do not know how

these factors specifically will play out in the basin

assessment area under climate-change scenarios,

but we can surmise some general possible outcomes for upper elevation communities.

Figure 34—Total species functional diversity (number of KEFs per community times the mean

number of species per function), by terrestrial vegetation community of the basin assessment

area (see table 2 for description of terrestrial vegetation communities).

57
Upper elevation vegetation communities, particularly alpine tundra and subalpine forest communities, may experience higher summer temperatures

and precipitation regimes. Under the “individual

response” hypothesis (Gleason 1926, Grime

1977), plant species would respond individually

and differently to climate changes (Covington and

others 1994, Mehringer, n.d.). Mehringer (1995,

n.d.) notes that upper elevation vegetation can

respond rapidly to climate change; spruce (Picea

spp.) and subalpine fir (Abies lasiocarpa) above

treeline in the Canadian Rocky Mountains

showed rapid upslope advances to >110 m above

present treeline between 9,000 and 5,000 years

ago and again near 1,000 years ago. Chapin and

others (1995) note that arctic tundra has responded to climate changes by altered nutrient

availability, growth-form composition, net

primary production, and species richness.

Alpine tundra and subalpine forest communities

currently are distributed among many mountain

ranges with differing degrees of geographic isolation (fig. 35). At present in the basin assessment

area, alpine tundra covers 95 100 ha (83 percent

of this occurs on BLM or FS lands and 17 percent

on other lands), and subalpine vegetation communities cover 6 633 600 ha (94 percent of this

occurs on BLM or FS lands and 6 percent on

other lands). Upward elevation shifts of these

communities may result in increasing isolation of

closely associated plant and animal species, reduction in overall distribution and area of occurrence in the basin assessment area, and local elimination of these communities and closely associated species. Mehringer (n.d.) notes that, because

of great topographic relief in the Columbia River

watershed, even slight changes of temperature

regimes and vegetation response can have a great

influence on total areas of vegetation types.

Some of the hot spots of biodiversity and species

rarity and endemism identified in Marcot and

others (1997) occur on isolated mountain tops,

and thus may be at risk from rapid climate

changes. The degree of risk depends on how

quickly the climate changes, how variable the

climate becomes during the change period, and

how well individual species can respond to such

variability either through genetic plasticity or by

changing aspect, elevation, or topographic location, as was found for Mojave species (Spaulding

and others 1983). Topographic situation—such

as mountain tops or upper elevation zones—may

not be limiting if changes occur slowly enough to

permit plant species to respond. One example of

a hot spot of biodiversity in upper elevation or

mountain settings is the eastern edge of the northern Cascade Range of Washington. In this area,

glaciated landscapes, rounded peaks, high elevations, and subalpine climates provide for unique

assemblages of amphibians and large carnivores,

in contrast with the surrounding landscapes, and

hold wetlands, bogs, and meadows that house

many rare plants. Another biodiversity hot spot

in mountain top settings is Glacier National Park

and the higher elevations of Flathead National

Forest in northwestern Montana, where rare large

carnivores and intact rare plant communities

occur in relatively undisturbed and isolated subalpine and alpine Rocky Mountain environments.

Three examples of hot spots of species rarity and

endemism in mountain top settings in Oregon

include the crest of the Umatilla Mountains,

where many west-side species are isolated and

disjunct on these mountain tops; Wallowa Mountain peaks and Sacajewea peak, which hold disjunct Rocky Mountain and boreal species; and

the Three Sisters area in Deschutes National Forest, which has unique volcanic substrates and

geologic landforms that house locally endemic

plants. Other examples are listed in Marcot and

others (1997).

Other ecological aspects of upper elevation communities may be affected by shifts in precipita9 Covington and others (1994:45) report that “general

circulation models (GCM) predict an increase of 1.5-2.5o C

in mean annual temperature within the next 30 to 50 years....

Actual meteorological records for the 1980 decade

and for the century indicate the Inland West region is

particularly vulnerable to global warming and to extreme

moisture stress.” Results of the regional climate model

simulations conducted for the present study, however,

suggest a more complex picture for the basin assessment

area (see footnote 2, chapter 3). The difference between the

model predictions may be because of the difference in

spatial resolution of the models. Covington and others do

concur with transition conditions to a new, future

equilibrium of (debatably) higher temperatures being

marked by extreme climatic variability and discontinuities.

Obviously, the final word on climate modeling and regional

effects has not been written, and much work remains to be

done, particularly in reconciling general circulation climate

models with regional climate models.

58
tion, temperature, or disturbance (especially fire)

regimes (Tausch and others 1993). As compared

with lower elevation forests, subalpine forests

are characterized by colder annual temperatures,

heavy winter snowpacks, forest floors as important “sinks” of total ecosystem nitrogen (“loose”

nitrogen cycles), greater importance of mycorrhizae in nutrient uptake, slower growth rates of

trees, greater carbon allocation to root biomass,

long retention of foliage, and long lifespan of

trees (Vogt and others 1989), as well as high arboreal epiphyte loads and occurrence of some

closely associated plant and animal species, including endemics. Some of these characteristics,

particularly nitrogen sinks, retention of foliage,

and lifespan of trees, may help buffer otherwise

adverse community responses to quick changes

in environmental conditions, including climate

change. But other characteristics, particularly mycorrhizae importance, and occurrence of close

associates and endemics, may impede quick

“adaptation” to new climatic and disturbance regimes. Overall, potential changes in climate and

disturbance patterns, and potential community

response, are likely to be complex.

Local climate changes in lowland environments

also may result in changes in fire regimes. Plants

that respond differently to occurrence, intensity,

frequency, and seasonality of fires likely would

show differential increases or decreases under

changing fire regimes. Marcot and others (1997)

present a list of rare or potentially rare plants of

the basin assessment area showing their orientation to fire regimes; this list can be consulted to

hypothesize how species might respond. Results

are quite mixed. For example, overstory lethal

fires may benefit 7 lichen groups, 9 plant groups,

and 9 vascular plant species, but harm a different

set of 5 bryophyte groups, 11 lichen groups,

2 plant groups, and 11 vascular plant species. Increasing fire frequency, increasing firesuppression activities, and changes in the seasonality of prescribed and natural fires—all of

which may occur from regional changes in

climate—also have mixed results.

ICBEMP

Legend

Alpine

Subalpine

State boundaries

Basin boundary

Figure 35—Current distribution of alpine tundra and subalpine forest communities in the

basin assessment area.

59
Invertebrate responses to changes in fire regimes

is poorly studied and largely unknown, except

for forest insect pests (Hann and others 1997).

Among vertebrates, stand-replacing fires can certainly change community composition. Hutto

(1995) documented differential response by bird

species to stand-replacement fires in the northern

Rocky Mountains, and noted that one species—

black-backed woodpecker (Picoides arcticus)—

was nearly restricted to sites with standing firekilled trees. Hutto also listed other bird species

that may benefit over the long term from standreplacement fires.

In some settings, vertebrate species may show

clearer responses to changes in climatic conditions than would plants. We identified vertebrate

species closely associated with upper elevation

vegetation communities and thus possibly at

greatest risk from upward altitudinal shifts of

these life zones. In the basin assessment area,

some 53 species—5 amphibians, 1 reptile, 22

birds, and 25 mammals—are associated with

alpine tundra communities. Many of these species

also occur in other vegetation communities. Eight

of these species (5 birds and 3 mammals) are

closely associated with alpine tundra communities (that is, occur in <20 percent of all 44

vegetation cover types) (fig. 36) and thus may be

at greatest risk from warming trends. Of these

eight species, white-tailed ptarmigan (Lagopus

leucurus) is found solely in alpine tundra, and

American pipit (Anthus rubescens) occurs in only

one other vegetation cover type (that is, occurs in

<10 percent of all 44 vegetation cover types).

Ptarmigan and pipits are especially vulnerable to

changes in alpine tundra communities.

Subalpine forest communities are potentially

vulnerable to upper elevation warming trends

(Graumlich 1991, Romme and Turner 1991,

Schullery 1995), although Peterson (1995) reports

that subalpine forests have been increasing in the

West because of warmer climates. Peterson

Figure 36—Vertebrate species at risk of regional warming trends, through decreases in total area of alpine

tundra. Species shown are closely associated with alpine tundra vegetation communities (that is, occur in

alpine tundra and <20 percent of all 44 vegetation cover types) in the basin assessment area.

60
(1995) also concludes that growth response of

subalpine forests varies considerably according

to geographic climate settings and microsite conditions. In the basin assessment area, subalpine

forest communities include Engelmann spruce

(Picea engelmannii)/subalpine fir forest, mountain hemlock forest, whitebark pine forest, and

whitebark pine/subalpine larch forest cover types.

Whitebark pine forest may be particularly vulnerable to climatic warming. Schullery (1995) projected that existing whitebark pine in Yellowstone National Park could decline by about 90

percent under a modest increase in warmth and

dryness (also see Whitlock 1993).

A total of 205 vertebrate species—20 amphibians,

6 reptiles, 106 birds, and 73 mammals—are associated with subalpine forest communities. Of

these, 36 species (8 amphibians, 16 birds, and 12

mammals) are closely associated with subalpine

forest communities (that is, occur in <20 percent

of all 44 vegetation cover types) and thus may be

at particularly greater risk from upper elevation

climatic warming (fig. 37). Of these 36 species,

13 occur in few other vegetation types (<10 percent of all 44 vegetation cover types) and are

particularly vulnerable. Among these 13 highly

vulnerable species, the salamanders include the

regionally endemic Cascade torrent salamander

Figure 37—Vertebrate species at risk of regional warming trends, through decreases in total area of subalpine

forests. Species shown are closely associated with subalpine forest vegetation communities (that is, occur in

subalpine forests and <20 percent of all 44 vegetation cover types) in the basin assessment area.

61
and Dunn’s salamander, and the locally endemic

Larch Mountain salamander; the birds include

spruce grouse (Dendragapus canadensis), boreal

owl (Aegolius funereus), black scoter (Melanitta

nigra), surf scoter (M. perspicillata), whitewinged scoter (M. fusca), oldsquaw (Clangula

hyemalis), three-toed woodpecker (Picoides

tridactylus), and boreal chickadee (Parus

hudsonicus); and the mammals include blacktailed deer (Odocoileus hemionus columbianus)

and California myotis (Myotis californicus).

The three aforementioned regionally or locally

endemic salamanders may be the most vulnerable

of all 13 species in terms of changes of subalpine

forest communities within the basin assessment

area. The three aforementioned scoters occur in

the basin assessment area as occasional migrants;

the surf scoter is an occasional winter visitor,

although the white-winged scoter has been recorded nesting in northeastern Washington. There

is likely very little danger to the viability of any

of the overall populations of scoters caused by

habitat changes within the basin assessment area.

Black-tailed deer ranges mostly west of the

Cascade Range and occurs in the basin assessment area only along the high Cascade Range.

Climate change might reduce the area of suitable

habitat for black-tailed deer within the basin assessment area, although they likely would persist

west of the crest of the Cascade Range. The rest

of the subalpine forest community species listed

above occur more regularly or more extensively

within the basin assessment area.

We expect that increasing isolation of upper elevation environments might result in differential

local isolation or extinctions of some animals,

much as past rising temperatures likely drove

some mammal species and their habitats upslope

into disjunct distributions on mountaintop islands

in the Great Basin (Diamond 1984). Likely survivors would include small mammals over big

mammals of the same trophic status and habitat

preference, habitat generalists over habitat specialists, and habitat generalist herbivores over

similarly sized habitat generalist carnivores (Diamond 1984). In alpine tundra communities in the

basin assessment area, such likely survivors might

include the following herbivorous small mammals

(≤ squirrel size) that are habitat generalists (occur in ≥50 percent of all 44 vegetation cover

types): deer mouse (Peromyscus maniculatus),

Columbian ground squirrel (Sperm ophilus

columbianus), golden-mantled ground squirrel

(S. lateralis), long-tailed vole (Microtus

longicaudus), and masked shrew (Sorex

cinereus).

The converse of these criteria might describe

mammal species more ecologically at risk of

climate-induced changes in alpine tundra habitat.

These species include big mammals (> squirrel

size) over small mammals, habitat specialists

(<50 percent of all 44 vegetation cover types)

over habitat generalists, and habitat generalist

carnivores over similarly sized habitat generalist

herbivores. In the basin assessment area, the one

at-risk mammal of alpine tundra fitting these criteria is wolverine.

Neilson (1991, 1993) and Noble (1993) note that

ecotones between biomes may be sensitive areas

to climatic change and therefore useful for monitoring change. Specifically, two types of change,

boundary shifts of regions and physiognomic

shifts within regions, are potentially independent

because of differential species responses. Each

type of change may require different strategies

for monitoring.

The response of plant, invertebrate, and vertebrate communities to any directional changes in

climate regimes likely will be made by individual

species rather than by entire communities. In

some cases, however, coadapted species complexes may suffer greater disruption than individual species’ habitat associations may suggest.

These complexes may include insect-plant pollination relations, invertebrate phoresis (especially

dependence on other organisms for dispersal),

mychorrhizal fungi-tree relations, and other

mutualistic, commensal, and symbiotic relations.

For example, Clark’s nutcracker (Nucifraga

columbiana) is a key element in regeneration of

whitebark pine—on the decline in the assessment

area—by caching (burying) seeds in the ground.

Clark’s nutcracker occurs in 16 vegetation cover

62
types in the basin assessment area. It is unclear

how complex shifts in plant species, vegetation

cover types, and disturbance regimes resulting

from climate change might affect Clark’s nutcracker populations. Wells (1983) hypothesized

that Clark’s nutcracker (along with pinyon jay

(Gymnorhinus cyanocephalus)) might have facilitated brisk redistribution of pinyon pine (Pinus

edulis) in the intermountain West following rapid

Holocene (Recent) climate changes. However, if

local, high-elevation populations of Clark’s nutcrackers become reduced and isolated because of

warming-induced declines in total area of montane and subalpine forests (the species does not

typically occur in alpine tundra communities or

much below montane forest zones), there may be

a compounding adverse affect on whitebark pine

redistribution and viability (see Marcot and others

1997 for further discussions of other vertebrate

species relations).

Another relation that might complicate species

response to changing climatic and disturbance

regimes is that of mycorrhizal fungi, their host

tree species, and soils. Disturbances that reduce

aeration or soil organic matter reduce mycorrhizal

activity (Amaranthus and Perry 1994). Reduction

in mycorrhizae reduces growth of many species

of trees and retards productivity of soils. Some

studies suggest that sites with harsh, continental

climates (such as parts of the basin assessment

area) may have lower fungal diversity than

marine, coastal climates, and thus may be poorly

buffered against changes in fungus populations

caused by disturbance (Powers 1989) or by adverse climate change.

Climate change also has the potential to alter decomposition processes in grasslands and coniferous forests (Anderson 1991, Breymeyer and

Melillo 1991, Esser 1992), plant respiration

(Ryan 1991), phenology and growth of alpine

vegetation (Walker and others 1994, 1995), bog

formation (Foster and Wright 1990), and the role

of soil in carbon sequestration (Van Veen and

others 1991). Van Veen and others (1991) report

that an increased level of CO2 (a so-called

“greenhouse gas”) in the atmosphere likely will

result in an increased input of organic carbon

into the soil because of the expected increase in

primary production. Whether this will lead to

accumulation of greater amounts of organic

carbon in soil depends on the flow of carbon

through the plant into the soil, the tightness of the

carbon cycle, and its subsequent transformation in the soil by microorganisms.

In this section, we have discussed potential future

changes of terrestrial ecological communities and

processes resulting from climate shifts. In the next

section, we peer backwards and discuss prehistoric changes in climates and in the diversity of

biota.

Paleoecology and Trends in

Vertebrate Species and

Communities of the Basin

Assessment Area

Late Quaternary Conditions and

Changes of the Basin Assessment

Area

The Quaternary period (Pleistocene and Holocene

[Recent] epochs of 1.64 million years ago to the

present) represents a time of remarkably variable

climates and biota in the inland West (McDonald

1984, Robbins 1993, Robbins and Wolf 1994).

Climate cycles during this period were a major

force in the evolution of plant response to change,

so that, according to Tausch (1993), current plant

communities today may be far less stable than

they appear to be from our perspective.

Using pollen records from the late Quaternary,

Mehringer (1985) reconstructed the following

vegetation changes in selected sites of the inland

West. Pollen sequences from the Holocene suggest fluctuations in vegetation resulting from

short, acute climatic changes and from fire and

volcanic episodes. Continental ice, alpine glaciers, and vast pluvial lakes had covered much of

the area during the last glacial episode. Most of

the northern Great Basin and adjacent provinces

were covered by cold sagebrush-steppe conditions

with montane conifers growing at lower elevations than they occur today.

Then, before 9,000 years ago during the early

Holocene, a warming trend caused shrinking

lakes, wasting glaciers, and catastrophic flooding

(Allen and others 1986). Pioneer plants such as

buffalo-berry (Shepherdia canadensis (L.) Nutt.)

increased and forests spread northward, while

63
sagebrush-steppe spread southward as montane

forests retreated to upper elevations. Douglas-fir

began its spread to become a dominant element

in conifer forests (Hermann 1985).

By 7,000 years ago, conifer forests lost ground to

grasslands and sagebrush, and shadscale and sagebrush communities replaced grasslands. By 5,400

years ago, this trend slowed, and by 4,000 years

ago, it reversed. A moister maritime climate developed in northern Idaho by 2,500 years ago or

sooner, and fire regimes changed (Mehringer

1985). By the late Quaternary, lodgepole pine

had spread north from Pleistocene glacial refuges

farther south (Critchfield 1985). Relict lodgepole

pine populations persisted in the Rocky Mountains (MacDonald and Cwynar 1985). Eventually,

Douglas-fir became an important part of Northwestern forests during interglacial periods of the

Quaternary (Hermann 1985).

The Quaternary fauna—known mostly from

mammal records—was characterized by the

spread of many large mammals throughout the

inland West, as summarized by Potts and

Behrensmeyer (1992). These included genera

(1) now globally extinct, such as the mammoth

(Mammathus), the bovid Soergelia, and sabertoothed cat (Smilodon); (2) extinct from the basin

assessment area but persisting today elsewhere,

such as jaguar (Panthera); and (3) persisting

within the basin assessment area to this day, including wolf (Canis), mustelids (including ermine), moose (Alces), caribou (Rangifer), horse

(Equus), cricetid rodents of genus Clethrionomys,

hare (Lepus), and vole (Microtus). Five Quaternary families of artiodactyls occurred, including

bovids, camelids, and antilocaprids, as well as

xenarthrans, including glyptodonts, armadillos,

and ground sloths. Proboscideans included

gomphotheres and mastodonts in addition to

the mammoths. Cricetid rodents underwent

exceptional adaptive radiation during the PlioPleistocene as savanna and grassland ceded to

steppe environments.

During the Pleistocene, rodents diversified, while

other groups (proboscideans, perissodactyls,

xenarthrans, and camelids) reduced. Immigrants

across the Bering land bridge to the area during

this time included bison (Bison), mountain sheep

(Ovis), muskox (Ovibos), moose (Alces), lion

(Panthera), and humans (Homo sapiens) (Pileou

1991). Climatic fluctuations served to blend

biotas during this time, mixing these newcomer

immigrants with the existing resident sabertoothed cat, dire wolf (Canis), horse, mammoth,

many squirrels (sciurids), New World rodents

(cricetids), and bovids (Potts and Behrensmeyer

1992).

In the late Pleistocene, large mammals suffered

waves of extinctions between 15,000 and 9,000

years ago. This “megafaunal extinction” took out

mammoths, mastodonts, gomphotheres, sabertoothed cats, dire wolves, horses, sloths, camels,

and many species of bears, deer, antilocaprids,

and others. It may be important to understand the

reasons for these extinctions—as yet, they are not

fully understood—to better interpret potential

effects of our modern day changes in climates

and habitats. Causes of the megafaunal extinction

episodes have been variously attributed to human

hunting, climate changes and associated habitat

shifts during the early interglacial periods, interspecific competition, and differential resorting of

species into increasingly disjunct communities

(Martin and Klein 1984).

Glacial cycles in the Holocene caused many

mammalian taxa to separate in distribution as

habitats became more patchy and disjunct and

as mammal species responded individually to

changing environments (Potts and Behrensmeyer

1992). The resulting communities have been

called nonanalogue communities or disharmonious faunas. Such faunas had high beta diversity

of species among geographic locations, such as

different mountain ranges. Some authors attributed these changes and the persistence of nonanalogue communities to periods of more equable

climates (Potts and Behrensmeyer 1992).

Several important lessons for interpreting

present-day biota in the basin assessment area

can be learned from this brief review of Quaternary period changes over the span of a dozen

millennia:

1. Climate shifts have occurred unpredictably and

have signalled great changes in both distribution

and species composition of floras and faunas.

64
2. Episodic extinctions of at least large mammals

have occurred, perhaps associated with climate

change, human hunting pressure, or other factors.

3. Differential extinctions and originations

(speciation and immigration of existing species)

have occurred mostly with unpredictable patterns

but over periods of millennia. Examples are the

great reduction in artiodactyls (especially horses)

and the increase in diversity of cricetids (mice

and rats) and sciurids (squirrels and chipmunks).

4. Separation and isolation of species because of

climatic shifts resulted from differential response

and sorting by species during periods of climate

change and equilibration.

At least as a working hypothesis, we might conjecture that today’s biotas of the basin assessment

area might respond with such similar, general patterns to increasing environmental pressures,

changing habitats, and potentially shifting climates. Unknowns in this hypothesis include (a)

rates of response of species, assemblages, and

ecological communities, which may be unable to

adequately respond and thereby might vanish; (b)

species-specific changes that may ensue, although

we can now draw from databases on speciesenvironment relations to begin to frame specific

predictions (for example, see the section above

on potential near-future climate change); and (c)

how effects of human activities might compound

with background natural variations in climates

and environments in affecting species and ecological communities. In this vein, historic and current human activities of greatest concern on Federal lands include short-term changes of natural

disturbance regimes such as fire, and human

activities directly affecting populations or habitats including hunting, urbanization, livestock

grazing, mining, and timber harvesting.

Two still-unanswered questions about climate

change during the Quaternary within the basin

assessment area may prove of interest to

management:

1. An explicit test is needed of the null hypothesis

that the range of historic conditions (past 50 to

200 years) does not reflect the range of prehistoric (Holocene) conditions, in terms of climate

and vegetation. To test this hypothesis, one might

use pollen diagrams of vegetation change (for example, Mehringer 1985) as compared with the

projections of historic vegetation conditions

(Hann and others 1997).

2. An explicit test is needed of the null hypothesis that there is no long-term (Holocene or Quaternary) set of repeating conditions of climate and

vegetation patterns within which an idealized,

stable range of natural variability (sensu Morgan

and others 1994) can be estimated. This hypothesis can be tested by calculating a running SE of

climate parameters or vegetation conditions (as

taken from pollen profiles), as a function of time,

and determining if it reaches an asymptote, thus

signifying a stable range of natural variability.

The first question above is important in determining if the range of historic conditions truly provides for environmental conditions in which species immigrated or evolved, and in which communities developed. The second question is important for testing the very concept of range of

natural variability.

Assuming that understanding the past and present

helps us project the future, additional work on the

following ecological topics would prove useful:

1. Identify Quaternary source pools of species,

zones and directions of spread of species, and

species dispersal dynamics, rates, and barriers.

2. Identify evolutionary adaptations and selection

mechanisms during the Quaternary, including

allopatric speciation, character divergence among

sympatric species, isolating mechanisms, and species swamping.

3. Study reproductive ecology pertinent to understanding adaptations, including hybridization

(Samson and Knopf 1994).

4. Study development of keystone roles of species

and their expression in changing environments

and communities.

5. Identify changes in endemism by taxonomic

level and proportion of taxa, and mechanisms

causing endemism (such as Pleistocene refugia,

allopatric or sympatric speciation, and isolation

by physiographic conditions such as montane

islands, flood-carved gorges, and scabland

landscapes).

65
Tertiary Biota and Comparison With Extant

Vertebrates in Eastern Oregon

Work at John Day Fossil Beds National Monument has described the ancient (pre-Pliocene

epoch, Tertiary period) flora and fauna of eastern

Oregon (Retallack and others 1996).10 The beds

occur in several arms of the John Day River system in what is now mainly grazed rangeland, agriculture land, and some sagebrush-steppe habitat.

Fossil records from the beds span more than 40

million years and date from 45 to 5 million years

ago. Although a few amphibians, reptiles, and one

raptor have been identified in fossil records at the

beds, the most diverse fossil evidence is that of

medium- to large-sized mammals. At the beds,

some 12 orders, 47 families, 155 genera, and 298

species of mammals have been identified spanning the 40-million-year period (fig. 38; see

footnote 10).

Of these Tertiary taxa, 8 orders (67 percent),

16 families (34 percent), 5 genera (3 percent), and

no species (0 percent) remain extant in the basin

assessment area (fig. 39). The Tertiary mammal genera still exant from the Tertiary are two

carnivora, Martes (Mustelidae) and Canis

(Canidae); two rodentia, “Sciurus” and

Spermophilus (both Sciuridae); and a chiropteran

bat, Myotis (Vespertilionidae). In addition, still

extant in the basin assessment area are the

amphibian (frog) Rana (Ranidae), and the turtle

Clemmys (Testudinidae). The current species

found within the general basin assessment area representing these ancient forms are listed in table

9. These are taxa of particular paleoecological

interest because of the antiquity of the genera.

Fossils of all these ancient genera were taken

from formations representing the mid-late

Miocene (8 to 6 million years ago), the most

recent of the Tertiary formations in John Day

Fossil Beds. An exception was “Sciurus,” which

came from the much older formations spanning

the late Eocene, Oligocene, and early Miocene

(39 to 20 million years ago). In addition, Clemmys

and Martes also were found in formations from

the mid-Miocene (15 to 12 million years ago).

Many other ancient families are still extant in the

basin assessment area even though none of their

Tertiary genera has survived. These ancient

families, listed in table 10, include those given

above as well as additional families of turtles,

lizards, avian raptors, marsupials, carnivores,

horses, pronghorns, rodents, and rabbits.

In comparison, the total extant mammal fauna

of the basin assessment area consists of 8 orders,

24 families, 67 genera, and 132 species (fig. 38).

Compared to all mammalian fossil taxa discovered at John Day Fossil Beds, the number of

extant orders is 67 percent of the total number of

Tertiary orders; families are 51 percent; genera

are 43 percent, and species are 44 percent. In

terms of taxonomic richness (number of taxa),

then, the Tertiary mammal fauna was far richer

than that at present, at all taxonomic levels.

This comparison, however, is uneven in several

ways. (1) The fossil record differentially preserves evidence of species with various life forms,

body size, and habitat associations. For example,

bats are represented by only a single species in

the fossil records of the beds, whereas the current

bat fauna includes some 15 species; more than a

single species probably existed prehistorically,

but other species have not been preserved or

detected yet in the fossil record. As well, rare species probably often go undetected in fossil records

(Koch 1978). (2) The mammalian fossil records

at the beds represent an enormous span of time as

compared with today’s “snapshot,” which is taken

over a much broader geography of the basin assessment area. Results of the above comparisons

would be even more exaggerated if confined to

just the extant sagebrush-steppe fauna, because

the current local biota of just the sagebrush-steppe

environment of eastern Oregon is substantially

less diverse than that of the full basin assessment

area. (3) Taxonomic problems of comparison

include the fact that species are more difficult

toidentify in the fossil record than are orders or

families, and prehistoric morphospecies do not

exactly coincide with current species in terms

of equivalent systematics. Still, the differences

between prehistoric and extant number of taxa

10 Personal communication. 1996. T. Fremd, John Day

Fossil Beds National Monument, John Day, Oregon.

66
Figure 38—Biodiversity (number of taxa) of prehistoric (Tertiary) mammals of

John Day Fossil Beds, eastern Oregon; and extant (still living) mammals of the

basin assessment area. (Source of Tertiary biodiversity information: see

footnote 10 in chapter 4.)

Figure 39—Biodiversity of prehistoric (Tertiary) mammals of John Day Fossil

Beds, eastern Oregon, showing number and percentage of prehistoric taxa

persisting to the present.

67
Table 9—Currently extant species or subspecies of vertebrates occurring in the basin assessment

area that are represented by Tertiary genera discovered in John Day Fossil Beds National

Monument, eastern Oregon, including the formation in which each genus was discovereda b

Tertiary genus Extant species or subspecies Order, family Formationc

Rana R. aurora, red-legged frog Anura, Ranidae A

R. boylii, foothills yellow-legged frog

R. cascadae, Cascades frog

R. catesbeiana, bullfrog

R. clamitans, green frog

R. luteiventris, spotted frog

R. pipiens, northern leopard frog

R. sylvatica, wood frog

Clemmys C. marmorata, western pond turtle Chelonia, A, B

Testudinidae

Martes M. americana, American marten Carnivora, A, B

M. pennanti, fisher Mustelidae

Canis C. latrans, coyote Carnivora,

C. lupus, gray wolf Canidae A

“Sciurus” S. griseus, western gray squirrel Rodentia, C

S. niger, eastern fox squirrel Sciuridae

Spermophilus S. armatus, Uinta ground squirrel Rodentia, A

S. beecheyi, California ground squirrel Sciuridae

S. beldingi, Belding’s ground squirrel

S. brunneus, Idaho ground squirrel

S. columbianus, Columbian ground squirrel

S. elegans nevadensis, Wyoming ground squirrel

S. lateralis, golden-mantled ground squirrel

S. saturatus, Cascade golden-mantled ground squirrel

S. townsendii, Townsend’s ground squirrel

S. washingtoni, Washington ground squirrel

Myotis M. californicus, California myotis Chiroptera, A

M. ciliolabrum, western small-footed myotis Vespertilionidae

M. evotis, long-eared myotis

M. lucifugus, little brown myotis

M. thysanodes, fringed myotis

M. volans, long-legged myotis

M. yumanensis, Yuma myotis

a See footnote 10 in chapter 4.

b These living taxa belong to ancient genera and are of particular evolutionary and biogeographical interest. All species and

subspecies listed here are currently found within the basin assessment area although not necessarily at John Day Fossil Beds.

c Formations: A = mid to late Miocene, 8 to 6 million years ago; B = mid to Miocene, 15 to 12 million years ago; and C = late

Eocene, Oligocene, and early Miocene, 39 to 20 million years ago.

68
Table 10—Currently extant taxa of vertebrates occurring in the basin assessment area that are

represented by Tertiary families discovered in John Day Fossil Beds National Monument, eastern

Oregon, including the formation in which the family was discovereda b

Tertiary family

still extant Extant representative genera Order Formationc

Ranidae Rana—true frogs Anura A

Chelydridae

(=Emydidae, part) Chrysemys—turtles Chelonia D

Testudinidae

(=Emydidae, part) Clemmys—turtles Chelonia A, B

Boidae Charina—boas Squamata C

Falconidae Falco—falcons Falconiformes B

Didelphidae Didelphis—opossum Marsupalia C

Mustelidae Gulo—wolverine

Lutra—otter

Martes—marten, fisher

Mephitis—skunk

Mustela—ermine, weasels

Spilogale—skunk

Taxidea—badger

Procyonidae Procyon— raccoon Carnivora B

Canidae Canis—coyote, wolf Carnivora A, B, C

Urocyon—gray fox

Vulpes—kit fox, red fox

Ursidae Ursus—bears Carnivora A, C

Equidae Equus—horse Perissodactyla A, B, C, D

Antilocapridae Antilocapra—pronghorn Artiodactyla A, B

Aplodontidae Aplodontia—mountain beaver Rodentia C

Castoridae Castor—beaver Rodentia A, B, C

Cricetidae Clethrionomys—red-backed voles Rodentia A, C

(=Muridae, part) Lemmiscus—vole

Microtus—voles

Mus—mouse

Neotoma—woodrats

Ondatra—muskrat

Onychomys—grasshopper mouse

Peromyscus—mice

Phenacomys—vole

Rattusrats

Reithrodontomys—harvest mouse

Synaptomys—lemming

69
(fig. 39) suggest that the “turnover” rate (extirpation or extinction, as compared with formation or

invasion of new taxa in the area) has been increasing, from least to most, in orders, families, genera,

and species.

At the beds, mammalian diversity of taxa within

orders can be depicted. In the fossil record of the

Tertiary, several orders were particularly rich in

numbers of families, genera, and species. Listed

in sequence of decreasing number of component

taxa, they were Artiodactyla, Carnivora,

Rodentia, and Perissodactyla (fig. 40). Among

extant forms, the more diverse orders (in decreasing sequence) are Rodentia, Carnivora,

Chiroptera, Artiodactyla, and Insectivora (fig.

41). This comparison suggests major declines

in diversity of perissodactyls. Webb (1984) outlined declines in horses of North America, which

area major portion of the fossil record of perissodactyls of John Day Fossil Beds; and Gingerich

(1984) noted that some 56 percent of Wisconsinan

perissodactyls disappeared from the North American fauna without taxonomic replacement (although the general ecological function of herbivory has been continued with the appearance of

new cricetid and sciurid species). Artiodactyls,

carnivores, and rodents have always been diverse,

although consisting of different families, genera,

and species over geologic time. Insectivores and

Table 10—Currently extant taxa of vertebrates occurring in the basin assessment area that are

represented by Tertiary families discovered in John Day Fossil Beds National Monument, eastern

Oregon, including the formation in which the family was discovereda b (continued)

Tertiary family

still extant Extant representative genera Order Formationc

Geomyidae Thomomys—pocket gophers Rodentia B, C

Heteromyidae Dipodomys—kangaroo rats Rodentia B, C

Microdipodops—kangaroo rat

Perognatus—pocket mice

Sciuridae Ammospermophilus—antelope squirrel Rodentia A, B, C

Glaucomys—flying squirrel

Marmota—marmots

Sciurus—squirrels

Spermophilus—ground squirrels

Tamias—chipmunks

Tamiasciurus—forest squirrels

Leporidae Brachylagus—pygmy rabbit Lagomorpha A, B, C

Lepus—hare, jackrabbits

Sylvilagus—cottontails

Vespertilionidae Antrozous—bat Chiroptera A

Eptesicus—brown bat

Euderma—bat

Lasionycteris—bat

Lasiurus—bat

Myotis—myotis (bats)

Pipistrellus—pipistrelle (bat)

Plecotus—big-eared bat

a See footnote 10 in chapter 4.

b These ancient families are of particular evolutionary and biogeographical interest. Extant representative genera of these

ancient families are listed; most of these genera are not present in the Tertiary fossil record (see table 9).

c Formations: A = mid to late Miocene, 8 to 6 million years ago; B = mid Miocene, 15 to 12 million years ago; C = late Eocene,

Oligocene, and early Miocene, 39 to 20 million years ago; and D = early to mid Eocene, 54 to 37 million years ago.

70
Figure 40—Diversity of families, genera, and species among orders of

prehistoric (Tertiary) mammals of John Day Fossil Beds, eastern Oregon (see

text for sources).

Figure 41—Diversity of families, genera, and species among orders of extant

mammals of the basin assessment area. Note that the vertical axis is scaled the

same as in figure 40.

71
chiropterans may have increased in diversity or

may simply be underrepresented in the fossil

record. Overall diversity of mammalian families,

genera, and species among orders, calculated by

using the Shannon-Weiner diversity index, are

nearly identical in comparing Tertiary to current

faunas (fig. 42). Thus, overall mammalian diversity has remained more or less constant, although

taxonomic composition of the fauna—especially

below order level—has turned over.

Records from different formations of John Day

Fossil Beds also have allowed reconstruction of

several major climatic periods (Chaney 1956,

Retallack and others 1996; see footnote 10):

1. Fifty-four to 37 million years ago (Clarno Formation; early to mid Eocene epoch)—Tropical to

subtropical forests covered the area. Plants included hundreds of species, many new to science,

now preserved in fossils of seeds, nuts, fruits,

leaves, branches, stems, and roots. Mammals

included browsing bonototheres and amynodonts,

hyaenadonts, and Patriofelis predators.

2. Thirty-nine to 20 million years ago (John Day

Formation; late Eocene, Oligocene, early

Miocene epochs)—Environments were diverse in

the John Day Fossil Beds region, with deciduous

forests replacing the earlier subtropical forests.

Early Miocene flora was highly diverse. The

fauna included many mid- to large-sized mammals, such as dogs, cats, swine, oreodonts, horses,

camels, rhinoceroses, and rodents. Volcanic eruptions then occurred between 20 and 15 million

years ago.

3. Fifteen to 12 million years ago (Mascall Formation; mid Miocene epoch)—Highly fertile soils

of the region, along with a moderate climate, led

to development of grassland and mixedhardwood woodlands during this period. The

fauna included horses, camels, deer, bears,

weasels, dogs, and cats. Large mammals included

gomphotheres, rhinos, and bear-dogs.

4. Eight to 6 million years ago (Rattlesnake Formation; mid to late Miocene epoch)—During this

period, the climate may have become dryer and

cooler, and the environment dominated by grasslands. Evidence of horses, sloths, rhinos, camels,

peccaries, pronghorns, dogs, bears, and other species has been found.

This record by formation at John Day Fossil Beds

provides an opportunity to compare prehistoric to

current vertebrate faunas by general time period

(fig. 43). Numbers of taxa among families, genera, and species become increasingly disparate

across these formations and periods. The period

of 39 to 20 million years ago (John Day Formation) seems to harbor the greatest diversity in

known fossils of families and genera. Current diversity of families and genera of the basin assessment area does not match that of this time period,

and would even be far less if only current-day

mid- and large-bodied mammals (to match those

taxa more likely to persist and be discovered in

the fossil record) of sagebrush-steppe communities were considered.

Ecological Integrity of

Terrestrial Species and

Communities

In this section, we depict current ecological integrity of the basin assessment area for terrestrial

ecosystems. Karr (1990) urged that protection

of biodiversity be one aspect of protecting the

biological integrity of natural resource systems.

Several studies (for example, Karr 1991, Kerans

and Karr 1994) have used a broadly based, multiparameter index of biotic integrity (IBI) for measuring environmental quality of aquatic ecosystems and status of fish assemblages. Other aspects

72
Figure 42—Shannon-Weiner diversity of families, genera, and species

among orders of prehistoric (Tertiary) and extant mammals of John

Day Fossil Beds and the basin assessment area, respectively.

Figure 43—Number of orders, families, and genera of mammals by formation

(Tertiary period, John Day Fossil Beds, eastern Oregon) and at present (basin

assessment area). MYBP = million years before present.

73
of measuring ecological integrity, however, also

could pertain to long-term viability and evolutionary potential of species, and long-term environmental conditions. We expand on the IBI approach by including these additional components

in our evaluation of ecological integrity of terrestrial species and communities of the basin

assessment area.

Haynes and others (1996) assumed six possible

goals that may be of use for managing ecological

integrity of Federal lands in the basin assessment

area. Three of the six goals pertain to terrestrial

ecological conditions. They are consistent with,

but were not necessarily modeled after, goals

for maintaining ecological integrity as listed by

Grumbine (1994). We assumed that the three

goals may provide useful benchmarks for measuring progress for at least some of the terrestrial

components of ecological integrity. The three

goals are to maintain species viability, maintain

long-term evolutionary potential of species, and

manage for multiple ecological domains and

evolutionary timeframes.

In this analysis, we divided each of these three

goals into subcomponents. For each component,

we identified specific GIS themes representing

current conditions, as follows:

Ecological integrity goal 1—Maintain species

viability

Species components:

• Component (1a)—Distribution of threatened

and endangered vertebrate species

• Component (1b)—Distribution of threatened

and endangered plants

• Component (1c)—Distribution of locally

endemic vertebrate species

• Component (1e)—Distribution of rare plants

(as listed in Natural Heritage databases)

• Component (1f)—Distribution of candidate

category C1 and C2 vertebrate species as

listed by USDI Fish and Wildlife Service11

Habitat connectivity component:

• Component (1d)—Distribution of vertebrate

species occurring along key habitat corridors

Ecological integrity goal 2—Maintain long-term

evolutionary potential of species

Species component:

• Component (2a)—Distribution of disjunct

vertebrate species populations

Diversity components:

• Component (2b)—Mapped locations of biodiversity hot spots (plants and vertebrates

combined)

• Component (2c)—Mapped locations of species

rarity and endemism hot spots (plants and

vertebrates combined)

• Component (2d)—Mapped locations of centers

of concentration of biodiversity, and of species

rarity and endemism (plants and vertebrates

combined)

11 This chapter reports on analyses conducted through

October 1995. Since that time, USDI Fish and Wildlife

Service has published a change in their species status

program, essentially replacing the three candidate species

categories with a single category of candidates for listing

with a 1-year review period of this program change (Federal

Register, February 28, 1996, vol. 61, no. 40, p. 7596). In this

change, most of the species that were classified as category

2 or 3, and 303 taxa that were category 1 candidates, are no

longer included in the list of candidate species. Many plant

and animal species addressed in this assessment were

denoted as candidate category 1 or 2 when the data were

gathered. Of those 131 category 1 or 2 plants, 4 became

candidates in the Federal Register notice: Castilleja christii,

Erigeron basalticus, Sidalcea oregana var. calva, and

Thelypodium howellii spp. spectabilis. Of those 34 category

2 animals (none had been designated category 1 in the basin

assessment area), only mountain plover (Charadrius

montanus) and spotted frog (Rana luteiventris, prev. R.

pretiosa) were still designated as candidates. All other plants

and animals were dropped from the list of candidates. This

assessment chapter, however, retains the listings for two

reasons: (1) our analyses and data collection preceded the

ruling change; and (2) our charge was to address species’

ecologies and conservation status, and the category 2 status

in particular still helps to denote species of potential

conservation concern deserving attention in this charge.

74
Ecological integrity goal 3—Manage for

multiple ecological domains and evolutionary

timeframes

Peripheral species and unique vertebrate

assemblages by ERU:

• Component (3a)—Upper Klamath Basin

species

• Component (3b)—Owyhee Upland species

• Component (3c)—Northern Great Basin

species

• Component (3d)—Amphibians of the

Columbia Gorge

• Component (3e)—Snake River Headwaters

ERU and western Wyoming species

• Component (3f)—Upper Snake River ERU

species

Component (3g)—Western Montana species

Full trophic ladder component:

• Component (3h)—Vertebrate carnivores

For each component, we produced a Paradox®

database and an ArcInfo® GIS map theme to

show locations or distributions of conditions by

4th code HUC. Specific results are as follows,

along with further discussions of the ecological

foundation for each goal and its components.

Ecological Integrity Goal 1:

Maintain Species Viability

One aspect of ecological integrity pertains to the

viability of species. Several authors have developed various assessments of species components

for various ecosystems. Lyons and others (1995)

list 10 components of fish assemblages in westcentral Mexico that they judged representative of

ecological conditions of species and their habitats. The components they used included number

of native species, number of sensitive species,

and percentage of exotic species. Karr (1991) and

Kerans and Karr (1994) assessed water quality

integrity by measuring attributes of fish assemblages. In similar fashion, we identified a series

of mappable species and diversity components

for this first goal.

Species components—The following five integrity components relate to distribution of sets of

species of particular concern for maintaining

species viability.

Component (1a): Distribution of threatened and

endangered vertebrate species—Eight federally

listed threatened or endangered vertebrates occur

in the basin assessment area. Geographic information systems maps of species range distributions were available for seven (table 11). Distribution of marbled murrelet (Brachyramphus marmoratus) was not mapped; habitat for this species

occurs at the extreme periphery of its range within

the basin assessment area in the Cascade Range

in northern Washington. The database and map

constructed for this component display the number of threatened and endangered vertebrate species by 4th code HUC (fig. 44). More threatened

and endangered vertebrate species are located in

the Cascade Range in northern Washington, the

northern glaciated mountains of northern Idaho

and western Montana, central Idaho mountains,

and the Greater Yellowstone Ecosystem than

elsewhere throughout the basin assessment area.

Component (1b): Distribution of threatened

and endangered plants—Four federally listed

threatened or endangered plants occur within the

basin assessment area (table 12). Range locations

(point locations of site occurrences) were taken

from state Natural Heritage Program databases.

The database and map constructed for this component display number of threatened and endangered plant species by 4th code HUC (fig. 45).

Threatened and endangered plants occur scattered

singly in six general locations in the basin assessment area: the Washington-Idaho border, the

Hell’s Canyon area of Oregon and Idaho, western

Montana, southern Idaho, south-central Oregon,

and the Klamath Basin portion of Oregon.

Component (1c): Distribution of locally endemic

vertebrates—Some seven vertebrate taxa (species

or subspecies) are locally endemic within the basin assessment area (table 13). The database and

map for these species display percentage of the

area of each 4th code HUC with coverage by any

or all of these species (fig. 46). Most locally endemic vertebrates occur in eastern Washington,

central Idaho mountains, and most of the southern

tier of the basin assessment area through southern

Oregon, southern Idaho, and adjacent states.

Component (1e): Distribution of rare plants (as

listed in Natural Heritage databases from the

75
Table 11—Threatened (T) or endangered (E) vertebrates of the basin

assessment area

Species

code Scientific name Common name Status

FALPER Falco peregrinus Peregrine falcon E

GRUAME Grus americana Whooping crane E

HALLEU Haliaeetus leucocephalus Bald eagle T

STROCA Strix occidentalis caurina Northern spotted owl T

BRAMAR Brachyramphus marmoratus Marbled murrelet T

RANTCA Rangifer tarandus caribou Woodland caribou E

URSARC Ursus arctos Grizzly bear T

CANLIR Canis lupus Gray wolf E

ICBEMP

Legend

Species = 1

Species = 2

Species = 3

Species = 4

4th field hydrologic

units

State boundaries

Basin boundary

Figure 44—Numbers of threatened or endangered vertebrate species by 4th-code hydrologic unit in the

basin assessment area (see table 11 for list of species). Asterisks refer to HUCs with wintering bald

eagles: * = at least four threatened or endangered vertebrate species including wintering bald eagles,

** = three threatened or endangered vertebrate species including wintering bald eagles.

76
ICBEMP

Legend

Species = 1

4th field hydrologic

units

State boundaries

Basin boundary

Figure 45—Numbers of threatened or endangered plant species by 4th code HUC in the basin

assessment area (see table 12 for list of species).

Table 12—Threatened (T) or endangered (E)

plants of the basin assessment areaa

Species

code Scientific name Status

ASTAPP Astragalus applegatei E

HOWAQU Howellia aquatilis T

MIRMAC Mirabilis macfarlanei E

STEMAL Stephanomeria malheurensis E

a Also, Ute ladies tresses (Spiranthes diluvialis), a federally listed

threatened species, was discovered within the basin assessment

area August 1996, too late to be included in the analyses in this

report.

77
Table 13—Locally endemic vertebrates of the basin assessment area

Species

code Scientific name Common name

DICATE Dicamptodon aterrimus Idaho giant salamander

MICPEK Microtus pensylvanicus kincaidi Potholes meadow vole

PLELAR Plethodon larselli Larch Mountain salamander

SPEWAS Spermophilus washingtoni Washington ground squirrel

THOTLI Thomomys talpoides limosus White salmon pocket gopher

TYMPHC Tympanuchus phasianellus columbianus Columbian sharp-tailed grouse

CENURO Centrocercus urophasianus Sage grouse

ICBEMP

Legend

Species = 0%

Species > 0 − 20%

Species > 20 − 40%

Species > 40 − 60%

Species > 60 − 80%

Species > 80 − 100%

4th field hydrologic

units

State boundaries

Basin boundary

Figure 46—Percentage of all 4th-code hydrologic units in the basin assessment area that include

mapped distributions of locally endemic vertebrate species (see table 13 for list of species).

78
Heritage Program of The Nature Conservancy)—

Plants listed in the Natural Heritage databases

were considered of potential conservation concern for this component. Some 129 rare plant species were included in the mapping of this component (table 14). The list was generated by (a)

compiling all Heritage Program reports of species

of all taxonomic groups from all locations and

states overlapping the basin assessment area, (b)

extracting only those reports of plant species

within the basin assessment area boundary, and

(c) cross-indexing the scientific names with those

identified by the Science Integration Team’s plant

expert panels, as rare, candidate, threatened, endangered, or of special conservation concern.

Data included are element occurrences of recent

past collections and current locations of plants;

thus, the map does not reflect just presence of

these species at the immediate present, but also

recent past occurrences as well. The database and

map for this component show the numbers of rare

plant species by 4th code HUC (fig. 47). The

greatest numbers of rare plants occur in eight

disjunct locations in the basin assessment area:

the Cascade Range in central Washington, the

southern Washington-Idaho border, the Blue

Mountains of northeast Oregon, the Hell’s Canyon region of the Oregon-Idaho border, central

Idaho mountains, Owyhee Uplands of Oregon

and Idaho, the Cascade Range of central Oregon,

and the Klamath Basin of Oregon.

Component (1f): Distribution of candidate category C1 and C2 vertebrate species—Some 39

vertebrate species with candidate listing status

occur in the basin assessment area (table 15). Distribution range maps were available for 28 of

these species.12 The database and map show

numbers of candidate vertebrate species by 4th

code HUC (fig. 48). Candidate vertebrate species

are more numerous along the Cascade Range in

central Washington, southeastern Washington

into Idaho, the Columbia Gorge, the Cascade

Range in central and southern Oregon, the

Klamath Basin, the high desert area along the

northeastern border of Oregon and Idaho, and the

Greater Yellow Ecosystem in northwestern

Wyoming and southeastern Idaho.

Habitat connectivity along the Cascade

Range—Another aspect of describing ecological

integrity is identifying broad-scale connectivity

of habitats for vertebrates. We assumed that connectivity may be useful for maintaining welldistributed populations and viability of vertebrates. In the basin assessment area, broad-scale

habitat corridors were identified by overlaying

range distributions of vertebrates species. We

identified potential broad-scale habitat corridors

as those 4th-code HUCs that meet the following

criteria: (1) the HUCs all contained the same

vegetation communities, such as montane forest;

(2) the HUCs all provided for distribution of

vertebrates that do not occur elsewhere throughout

the basin assessment area; and (3) more than two

contiguous HUCs pertain to the first two criteria

(thus, disjunct HUCs do not qualify, and conditions met for only one or two contiguous HUCs

constitute a corridor at a finer scale of resolution

than addressed in this broad-scale assessment).

12 At the time of this analysis, ranges for only 28 species

were available. Subsequently, ranges for all 39 have been

mapped and are available.

79
Table 14—Rare plants of the basin assessment

area as listed in Natural Heritage databases

Species

code Scientific name

ALLAAS Allium aaseae

ALLANC Allium anceps

ALLCON Allium constrictum

ALLDIC Allium dictuon

ALLMAD Allium madidum

ALLROB Allium robinsonii

AMSCAR Amsinckia carinata

ANTARC Antennaria arcuata

ANTARO Antennaria aromatica

ARAFEC Arabis fecunda

ASTJES Aster jessicae

ASTMOL Aster mollis

ASTANS Astragalus anserinus

ASTAPP Astragalus applegatei

ASTCOL Astragalus columbianus

ASTHOW Astragalus howellii

ASTMUL Astragalus mulfordiae

ASTONI Astragalus oniciformis

ASTPAY Astragalus paysonii

ASTPEC Astragalus peckii

ASTSCA Astragalus scaphoides

ASTSIN Astragalus sinuatus

ASTSTE Astragalus sterilis

ASTTEG Astragalus tegetarioides

ASTTYG Astragalus tyghensis

ASTYOW Astragalus yoder-williamsii

BALROS Balsamorhiza rosea

BOTASC Botrychium ascendens

BOTCRE Botrychium crenulatum

BOTLUN Botrychium lunaria

BOTPAR Botrychium paradoxum

BOTPED Botrychium pedunculosum

BOTPUM Botrychium pumicola

CALNIT Calochortus nitidus

Table 14—Rare plants of the basin assessment

area as listed in Natural Heritage databases

(continued)

Species

code Scientific name

CASCHL Castilleja chlorotica

CASCHR Castilleja christii

CASCRY Castilleja cryptantha

CASFRA Castilleja fraterna

CASRUB Castilleja rubida

CHACUS Chaenactis cusickii

COLMAZ Collomia mazama

COLREN Collomia renacta

CYMDAV Cymopterus davisii

CYMDOU Cymopterus douglassii

CYMNIV Cymopterus nivalis

CYPFAS Cypripedium fasciculatum

DELVIR Delphinium viridescens

DESTOR Descurainia torulosa

DOUIDA Douglasia idahoensis

DRATRI Draba trichocarpa

ERIBAS Erigeron basalticus

ERILAC Erigeron lackschewitzii

ERISAL Erigeron salmonensis

ERICHR Eriogonum chrysops

ERICRO Eriogonum crosbyae

ERICUS Eriogonum cusickii

ERIMEL Eriogonum meledonum

ERIPRO Eriogonum prociduum

GRAHET Gratiola heterosepala

GRIHOW Grindelia howellii

HACCRO Hackelia cronquistii

HACVEN Hackelia venusta

HAPINS Haplopappus insecticruris

HAPLIA Haplopappus liatriformis

HAPRAD Haplopappus radiatus

HOWAQU Howellia aquatilis

ILILON Iliamna longisepala

80
Table 14—Rare plants of the basin assessment

area as listed in Natural Heritage databases

(continued)

Species

code Scientific name

LEPDAV Lepidium davisii

LEPGLA Leptodactylon glabrum

LESHUM Lesquerella humilis

LESPAY Lesquerella paysonii

LOMERY Lomatium erythrocarpum

LOMGRE Lomatium greenmanii

LOMSUK Lomatium suksdorfii

LOMTUB Lomatium tuberosum

LUISER Luina serpentina

LUPCUS Lupinus cusickii

MECORE Meconella oregana

MENMOL Mentzelia mollis

MENPAC Mentzelia packardiae

MIMCLI Mimulus clivicola

MIMHYM Mimulus hymenophyllus

MIMJEP Mimulus jepsonii

MIMJUN Mimulus jungermannioides

MIMLAT Mimulus latidens

MIMPAT Mimulus patulus

MIMPUL Mimulus pulsiferae

MIMPYG Mimulus pygmaeus

MIMSUK Mimulus suksdorfii

MIMTRI Mimulus tricolor

MIRMAC Mirabilis macfarlanei

MUSLIN Musineon lineare

OENPSA Oenothera psammophila

ORYCON Oryzopsis contracta

ORYHEN Oryzopsis hendersonii

PAPPYG Papaver pygmaeum

PENBAR Penstemon barrettiae

PENCOM Penstemon compactus

PENGLA Penstemon glaucinus

PENIDA Penstemon idahoensis

Table 14—Rare plants of the basin

assessment area as listed in Natural Heritage

databases (continued)

Species

code Scientific name

PENJAN Penstemon janishiae

PENLEM Penstemon lemhiensis

PENPEC Penstemon peckii

PERERY Perideridia erythrorhiza

PHAINC Phacelia inconspicua

PHALEN Phacelia lenta

PHAMIN Phacelia minutissima

PHLIDA Phlox idahonis

PLEORE Pleuropogon oregonus

POLPEC Polemonium pectinatum

POTCOT Potentilla cottamii

PRIALC Primula alcalina

RANREC Ranunculus reconditus

RORCOL Rorippa columbiae

RUBBAR Rubus bartonianus

RUBNIG Rubus nigerrimus

SENERT Senecio ertterae

SILSEE Silene seelyi

SILSPA Silene spaldingii

SISSAR Sisyrinchium sarmentosum

STEMAL Stephanomeria malheurensis

TAUHOO Tauschia hooveri

TEXSAJ Texosporium sancti-jacobi

THEEUC Thelypodium eucosmum

THEREP Thelypodium repandum

TRIDOU Trifolium douglasii

TRILEI Trifolium leibergii

TRIOWY Trifolium owyheense

TRITHO Trifolium thompsonii

81
ICBEMP

Legend

No species present

1−25 species present

26−50 species present

51−75 species present

76−100 species present

100+ species present

4th field hydrologic

units

State boundaries

Basin boundary

Figure 47—Numbers of rare plants (as identified in Natural Heritage databases from the Heritage Program or similar stateprograms of The Nature Conservancy) by 4th-code hydrologic unit in the basin assessment area (see table 17 for list ofspecies).

82
Table 15—Federally listed candidate category C1 or C2 vertebrates of the basin assessment areaa

Species

code Scientific name Common name

ACCGEN Accipiter gentilis Northern goshawk\a

AGETRI Agelaius tricolor Tricolored blackbird

ASCTRU Ascaphus truei Tailed frog

ATHCUN Athene cunicularia Burrowing owl

BRAIDA Brachylagus idahoensis Pygmy rabbit

BUFBOR Bufo boreas Western toad

BUTREG Buteo regalis Ferruginous hawk

CENURO Centrocercus urophasianus Sage grouse

CHLNIG Chlidonias niger Black tern

CLEMAR Clemmys marmorata Western pond turtle\b

CONBOR Contopus borealis Olive-sided flycatcher

CYGBUC Cygnus buccinator Trumpeter swan

EMPTRA Empidonax traillii Willow flycatcher\a

EUDMAC Euderma maculatum Spotted bat

GULGUL Gulo gulo Wolverine\b

HISHIS Histrionicus histrionicus Harlequin duck

IXOEXH Ixobrychus exilis hesperis Western least bittern

LANLUD Lanius ludovicianus Loggerhead shrike

LYNLYN Lynx lynx Lynx\b

MARPEN Martes pennanti Fisher\b

MICPKI Microtus pennsylvanicus kincaidi Potholes meadow vole

MYOCIL Myotis ciliolabrum Western small-footed myotis\a

MYOEVO Myotis evotis Long-eared myotis\a

MYOTHY Myotis thysanodes Fringed myotis

MYOVOL Myotis volans Long-legged myotis\a

MYOYUM Myotis yumanensis Yuma myotis

OVICCA Ovis canadensis californiana California bighorn sheep\b

PLECHI Plegadis chihi White-faced ibis\a

PLELAR Plethodon larselli Larch mountain salamander

PLETOW Plecotus townsendii pallescens Pale western big-eared bat

RANAUR Rana aurora Red-legged frog

RANCAS Rana cascadae Cascades frog

RANLUT Rana luteiventris Spotted frog

SCEGRA Sceloporus graciosus graciosus Sagebrush lizard

SORPRE Sorex preblei Preble’s shrew

SPEBRE Spermophilus brunneus Idaho ground squirrel

SPEWAS Spermophilus washingtoni Washington ground squirrel

TYMPHC Tympanuchus phasianellus columbianus Columbian sharp-tailed grouse

a See footnote 11 chapter 4 for recent changes in candidate species listings by USDI Fish and Wildlife Service. At the time of

this analysis, only 28 species’ ranges were available. (Subsequently, all 39 species’ ranges were mapped and are available.)

The remaining 11 species are marked as \a -scarce but ubiquitous or widespread in the basin assessment area or \b - not

ubiquitous but for which no range map was available for the entire basin assessment area.

83
ICBEMP

Legend

Species = 8 − 10

Species > 10 − 13

Species > 13 − 16

Species > 16 − 19

4th field hydrologic

units

State boundaries

Basin boundary

Figure 48—Numbers of federally listed candidate category C1 or C2 vertebrate species by 4th-code hydrologic unit in thebasin assessment area (see table 15 for list of species, and text footnote 11 for explanation of recent changes in thecandidate species listing).

84
Although some potential vertebrate habitat corridors could be identified—such as linking Glacier

National Park with Yellowstone National Park,

or the Blue Mountains and the central Idaho

mountains, for many montane or alpine species

including mustelids—they did not meet the second criterion for this particular component of

being the only locations of associated vertebrate

species within the basin assessment area. They

are included, however, in the “Full Trophic Ladder Component,” 3(h), discussed below. The following integrity component relates to habitat connectivity as defined above.

Component (1d):13 Distribution of species occurring along key habitat corridors—One major

habitat corridor that met all the above criteria

was found along the eastern spine of the Cascade

Range. An assemblage of seven vertebrates14

(table 16) occurs in the basin assessment area

only along the spine or eastern flanks of the

Cascade Range in Washington and Oregon collectively delineate a mid- to high-elevation montane corridor. The database and map show presence of this corridor by 4th code HUC (fig. 49).

The corridor occurs along the eastern flanks of

the Cascade Range; additional 4th code HUCs in

eastern Washington, northern Idaho, and western

Montana also contain disjunct populations of

these species and are indicated on the map.

Ecological Integrity Goal 2:

Maintain Long-Term Evolutionary

Potential of Species

The overall concept of maintaining long-term

evolutionary potential of species entails, in part,

maintaining unique ranges and centers of biodiversity, rarity, and endemism.

Species components—The following integrity

component relates to distribution of species that

may be critical to maintaining long-term evolutionary potential. Other species components to

help meet this goal include maintaining unique

ecotypes of species, maintaining species at the

periphery of their global range, and maintaining

unique subspecies, varieties, or phenotypes

(Marcot and others 1996). These additional components were not available for mapping in this

analysis.

Component (2a): Distribution of disjunct vertebrate species populations—Managing for disjunct populations of plants and animals is an

important facet of maintaining genetic diversity

and longterm evolutionary potential of species

(Kruckeberg and Rabinowitz 1985, Millar and

Libby 1991).

Some 11 vertebrate species are coded in the SER

database as having disjunct ranges within the basin assessment area. Two of these species, however, Virginia opossum (Didelphis virginiana)

and bullfrog (Rana catesbeiana), are mainly recent invaders or introduced, are not of conservation concern, and are not included in this analysis.

In fact, bullfrogs, widely introduced to ponds and

lakes throughout the basin assessment area, are

predators on some of the more increasingly scarce

native amphibians such as red-legged frog. A

third species with a disjunct range, pygmy shrew

(Sorex hoyi), is largely contiguous within the basin assessment area (as mapped at the broad scale

used by the expert panels for this project) with

only a small portion of its range disjunct, so it

13 This component is numbered out of sequence here to

remain consistent with the map index codes on record for

ICBEMP.

14 An additional, eighth species fitting these criteria is

mountain beaver (Aplodontia rufa), which was not mapped

in time for this analysis. Its inclusion would not alter the

results, as its distributional range within the basin assessment area matches that of the other species listed for this

component. Other vertebrate species such as northwestern

salamander, Cope’s giant salamander, ensatina (Ensatina

eschscholtzii), Larch Mountain salamander, Columbian

black-tailed deer (Odocoileus hemionus columbianus), and

others also occur within the basin assessment area only in

the Cascade Range, but not broadly along the spine or

eastern front.

85
Table 16—Vertebrates that occur within the basin assessment area only along a broad latitudinal

extent of the eastern flanks of the Cascade Rangea

Species

code Scientific name Common name

RANCAS Rana cascadae Cascades frog

TARGRA Taricha granulosa Rough skinned newt

STROCA Strix occidentalis caurina Northern spotted owl

SORTRO Sorex trowbridgii Trowbridge’s shrew

NEUGIB Neurotrichus gibsii Shrew mole

SPESAT Spermophilus saturatus Cascade golden mantled ground squirrel

ZAPTRI Zapus trinotatus Pacific jumping mouse

a These include species that might benefit by latitudinal habitat corridors of montane and subalpine forests along the eastern

slope of the Cascade Range. This list does not include species that enter the basin assessment area from the west only at one

point across the spine of the Cascade Range or only up the Columbia River Gorge.

ICBEMP

Legend

Present

Absent

4th field hydrologic

units

State boundaries

Basin boundary

Figure 49—Occurrence of a habitat corridor for vertebrates of the Cascade Range by 4th-code

hydrologic unit in the basin assessment area (see text for further explanation of the extraneous

hydrologic units shown shaded here).

86
was excluded from subsequent analyses. The remaining eight species with disjunct ranges within

the assessment area are included in the analyses

(table 17). Note that some of these eight species

also may occur in one of the components listed

above (for example, whooping crane (Grus

americana) is also an endangered species). Wild

turkey (Meleagris gallopavo) was included in this

list because, although its disjunct populations result from introductions, the species has naturalized in these ranges (it is a “desired nonnative

vertebrate” in the parlance of National Forest

Management Act Regulations 38 CFR 219) and

is a sought-after species of positive economic

and social importance. The database and map for

this component show percentage of the area of

each 4th code HUC with coverage by any or all

of these species (fig. 50). A count of species by

4th code HUC was not used as the measure

because, interestingly, many of these species are

distributed as allopatric (nonoverlapping ranges),

so that a simple count of species by HUC was

not a sensitive measure of distribution. The mapped results suggest that HUCs important to disjunct vertebrate populations occur in Montana,

northern Idaho, central Idaho mountains, Snake

River plain in southwest Idaho, along the

Washington-Oregon border, and in the Cascade

Range in Oregon eastward into the high desert

and Klamath Basin.

Diversity components—Component (2b): Mapped locations of biodiversity hot spots (of plants

and animals combined)—This theme was derived

by overlaying the maps of centers of concentration of biodiversity of plants and animals (component 2d, below) and generating polygons where

at least three centers overlap (see Marcot and

others 1997 for further information on conditions

within each hot spot). The database and map for

this component show percentage of the area of

each 4th code HUC with coverage by any or all

polygons of biodiversity hot spots (fig. 51). Seven

biodiversity hot spots were identified in various

locations throughout the basin assessment area.

Component (2c): Mapped locations of species

rarity and endemism hot spots (of plants and

animals combined)—This theme was derived by

overlaying the maps of centers of concentration

of species rarity and endemism of plants and

animals (component 2d, below), and generating

polygons where at least three centers overlap (see

Marcot and others 1997 for further information

on conditions within each hot spot). The database

and map show percentage of the area of each 4th

code HUC with coverage by any or all polygons

(fig. 52). Twelve hot spots of species rarity and

endemism were identified in various locations,

mostly in southern Washington, in Oregon, and

southwestern Idaho. One major concentration

area is the Columbia River Gorge.

Component (2d): Mapped locations of centers

of concentration of biodiversity, and of species

rarity and endemism (for plants and animals

combined)—This theme was derived as polygons

mapped by expert panels during the panel meeting

on natural areas (see Marcot and others 1997 for

further information on methods). The database

and map show percentage of the area of each 4th

code HUC with coverage by any or all polygons

(fig. 53). Collectively, centers of concentration

for plants and animals occur in many locations

throughout the basin assessment area.

Ecological integrity goal 3: manage

for multiple ecological domains and

evolutionary timeframes

The overall concept of managing for multiple

ecological domains and evolutionary timeframes

entails ensuring that unique species assemblages

by geographic area are recognized and provided

for, and that the full trophic ladder occurs (represented here by distributions of large vertebrate

carnivores). Other components of this goal also

may entail ensuring that the full range of historic

and current habitat conditions and terrestrial communities are provided, including mid-scale and

site-specific microhabitats and substrates; however, these components were not mapped for this

analysis.

87
Table 17—Examples of vertebrate species with disjunct distributions in the basin

assessment area, as mapped at the broad scale

Species

code Scientific name Common name

BUFWOO Bufo woodhousii Woodhouse’s toad

GRUAME Grus ameriana Whooping crane

LAMZON Lampropeltis zonata California mountain kingsnake

MELGAL Meleagris gallopavo Wild turkey

RANCLA Rana clamitans Green frog

RANSYL Rana sylvatica Wood frog

RHILEC Rhinocheilus lecontei Longnose snake

SONSEM Sonora semiannulata Ground snake

ICBEMP

Legend

Species = 0%

Species 1 − 20%

Species > 20 − 40%

Species > 40 − 60%

Species > 60 − 80%

Species > 80 − 100%

4th field hydrologic

units

State boundaries

Basin boundary

Figure 50—Percentage of each 4th-code hydrologic unit in the basin assessment area that includes

mapped distributions of vertebrate species with disjunct populations (see table 17 for list of species).

88
ICBEMP

Legend

Species = 0%

Species 1 − 20%

Species > 20 − 40%

Species > 40 − 60%

Species > 60 − 80%

Species > 80 − 100%

Biodiversity hot spots

4th field hydrologic

units

State boundaries

Basin boundary

Figure 51—Percentage of each 4th-code hydrologic unit in the basin assessment area that includes biodiversity hot spots ofplants and animals (source: Marcot and others 1997).

89
ICBEMP

Legend

Species = 0%

Species 1 − 20%

Species > 20 − 40%

Species > 40 − 60%

Species > 60 − 80%

Species > 80 − 100%

Species rarity and

endemism hot spots

4th field hydrologic

units

State boundaries

Basin boundary

Figure 52—Percentage of each 4th-code hydrologic unit in the basin assessment area that includes species rarity andendemism hot spots of plants and animals (source: Marcot and others 1997).

90
ICBEMP

Legend

NOTE:

Centers of biodiversity,

rarity and endemism are

outside of biodiversity

and endemism hot spots

Species = 0%

Species 1 − 20%

Species > 20 − 40%

Species > 40 − 60%

Species > 60 − 80%

Species > 80 − 100%

Centers of biodiversity,

rarity and endemism

4th field hydrologic

units

State boundaries

Basin boundary

Figure 53—Percentage of each 4th-code hydrologic unit in the basin assessment area that includes centers of concentrationof biodiversity, or of species rarity and endemism, of plants and animals (source: Marcot and others 1997).

91
Unique species assemblage components—The

following integrity components relate to identifying unique vertebrate species assemblages by

geographic area (ERU), as identified and listed

in Marcot and others (1997).15 Many of these

assemblages are of species that occur peripherally

in the basin assessment area. Managing for the

periphery of ranges of species can be important

for maintaining long-term evolutionary potential

(Furlow and Armijo-Prewitt 1995, Kruckeberg

and Rabinowitz 1985, Lesica and Allendorf

1995), for it is at the edges of ranges where environmental factors often differ the most and new

genotypes and phenotypes emerge (see Marcot

and others 1997 for examples in the basin assessment area).

Component (3a): Upper Klamath Basin species—

We identified a set of 10 species that are closely

associated with the Upper Klamath Basin ERU in

southern Oregon (fig. 54). The database and map

show number of species by 4th code HUC.

Component (3b): Owyhee Upland species—We

identified a set of nine species that are closely associated with the Owyhee Upland ERU in southern Oregon and Idaho (fig. 55). The database and

map show number of species by 4th code HUC.

Component (3c): Northern Great Basin species—

We identified a set of three species that are

closely associated with the northern Great Basin

ERU in southern Oregon (fig. 56). The database

and map show number of species by 4th code

HUC.

Component (3d): Amphibians of the Columbia

River Gorge—This unique assemblage consists

of amphibian species that enter into the basin

assessment area mainly or solely through the

Columbia River Gorge (fig. 57). The database and

map show number of species by 4th code HUC.

Component (3e): Snake River Headwaters ERU

and western Wyoming species—A set of eight

species were identified that are closely associated

with the Snake River Headwaters ERU in southeastern Idaho, and the western Wyoming (Greater

Yellowstone Ecosystem) region (fig. 58). The

database and map show number of species by 4th

code HUC.

Component (3f): Upper Snake River ERU

species—Two species were identified that are

closely associated with the Upper Snake River

ERU in southern Idaho (fig. 59). The database

and map show number of species by 4th code

HUC.

Component (3g): Western Montana species—

Two species were identified that are closely

associated with western Montana (fig. 60). The

database and map show number of species by 4th

code HUC.

Full trophic ladder component—Component

(3h): Vertebrate carnivores—Eight wide-ranging

vertebrate carnivores occur in the basin assessment area: grizzly bear (Ursus arctos), black bear

(U. americanus), gray wolf (Canis lupus), cougar

(Felis concolor), lynx (Lynx lynx), wolverine,

fisher (Martes pennanti), and American marten

(M. americana). Four of these species were

mapped and are included in this component (fig.

61).16 The full set of vertebrate carnivores—and

by inference, the full trophic ladder—once likely

ranged throughout the entire basin assessment

area, but now is present only in the northern

Rocky Mountains, including Glacier National

Park, Yellowstone National Park and adjacent

lands, and in other locations where grizzly bear

and gray wolf recovery areas overlap, as in central

Idaho and north-central Washington. The database and map for this component show number

of species by 4th code HUC (fig. 61).

Text continued on page 101

15 Since the current analysis was conducted, some range

maps of species have been refined, ranges of additional

vertebrate species of the basin assessment area have been

mapped, and additional species have been added to these

lists of unique species assemblages by ERU and geographic

area. The lists presented in Marcot and others (1997) reflect

these additions but were not included in time to update

analyses presented here. We do not anticipate, however, that

these additions would qualitatively change results presented

here.

16 Since this analysis, ranges for the remaining four carnivores also were mapped. Their distributional ranges more

than overlap those of the four initially mapped carnivore

species. Thus, their inclusion in this analysis does not change

the results of identifying centers of high-carnivore richness.

92
ICBEMP

Legend

Species = 0

Species = 1

Species = 2

Species = 3

Species = 4

Species = 8

Species = 10

4th field hydrologic

units

State boundaries

Basin boundary

Figure 54—Numbers of species closely associated with the Upper Klamath Basin ecology reporting unit (ERU), by 4th-codehydrologic unit in the basin assessment area (see Marcot and others 1997 for list of species).

93
ICBEMP

Legend

Species = 0

Species = 1

Species = 2

Species = 3

Species = 4

Species = 5

Species = 6

Species = 7

Species = 8

4th field hydrologic

units

State boundaries

Basin boundary

4th field hydrologic

units for

species = 8 only

Figure 55—Numbers of species closely associated with the Owyhee Uplands ERU, by 4th-code hydrologic unit in the basinassessment area (see Marcot and others 1997 for list of species).

94
ICBEMP

Legend

Species = 0

Species = 1

Species = 2

Species = 3

4th field hydrologic

units

State boundaries

Basin boundary

Figure 56—Numbers of species closely associated with the Northern Great Basin ERU, by 4th-code hydrologic unit in thebasin assessment area (see Marcot and others 1997 for list of species).

95
ICBEMP

Legend

Species = 0

Species = 1

Species = 2

Species = 4

4th field hydrologic

units

State boundaries

Basin boundary

Figure 57—Numbers of amphibian species closely associated with the Columbia River Gorge, by 4th-code hydrologic unit inthe basin assessment area (see Marcot and others 1997 for list of species).

96
ICBEMP

Legend

Species = 0

Species = 1

Species = 2

Species = 4

Species = 5

Species = 6

Species = 8

4th field hydrologic

units

State boundaries

Basin boundary

Figure 58—Numbers of species closely associated with the Snake River Headwaters ERU and western Wyoming, by4th-code hydrologic unit in the basin assessment area (see Marcot and others 1997 for list of species.)

97
ICBEMP

Legend

Species = 0

Species = 1

Species = 2

4th field hydrologic

units

State boundaries

Basin boundary

Figure 59—Numbers of species closely associated with the Upper Snake River ERU, by 4th-code hydrologic unit in the basinassessment area (see Marcot and others 1997 for list of species.)

98
ICBEMP

Legend

Species = 0

Species = 1

Species = 2

Species = 3

4th field hydrologic

units

State boundaries

Basin boundary

Figure 56—Numbers of species closely associated with the Northern Great Basin ERU, by 4th-code hydrologic unit in thebasin assessment area (see Marcot and others 1997 for list of species).

99
ICBEMP

Legend

Species = 0

Species = 1

Species = 2

Species = 4

4th field hydrologic

units

State boundaries

Basin boundary

Figure 57—Numbers of amphibian species closely associated with the Columbia River Gorge, by 4th-code hydrologic unit inthe basin assessment area (see Marcot and others 1997 for list of species).

100
ICBEMP

Legend

Species = 0

Species = 1

Species = 2

Species = 4

Species = 5

Species = 6

Species = 8

4th field hydrologic

units

State boundaries

Basin boundary

Figure 58—Numbers of species closely associated with the Snake River Headwaters ERU and western Wyoming, by4th-code hydrologic unit in the basin assessment area (see Marcot and others 1997 for list of species.)

101
ICBEMP

Legend

Species = 0

Species = 1

Species = 2

4th field hydrologic

units

State boundaries

Basin boundary

Figure 59—Numbers of species closely associated with the Upper Snake River ERU, by 4th-code hydrologic unit in the basinassessment area (see Marcot and others 1997 for list of species.)

102
ICBEMP

Legend

Species = 0

Species = 1

Species = 2

4th field hydrologic

units

State boundaries

Basin boundary

Figure 60—Numbers of species closely associated with western Montana, by 4th-code hydrologic unit in the basinassessment area (see Marcot and others 1997 for list of species.)

103
ICBEMP

Legend

Species = 2

Species = 3

Species = 4

4th field hydrologic

units

State boundaries

Basin boundary

Figure 61—Numbers of wide-ranging vertebrate carnivores (gray wolf, black bear, cougar, and coyote) by 4th-codehydrologic unit in the basin assessment area (see text and Marcot and others 1997 for full list of species.) * = include graywolf (not initially mapped).

104
Pictures of Terrestrial Integrity:

Combining Maps for Each Goal

The final step in depicting terrestrial ecological

integrity of the basin assessment area was to combine all components under each goal into maps

depicting overall conditions for each of the three

goals. Similarly, Lyons and others (1995) produced composite maps to depict overall integrity

conditions of fish assemblages in their study.

They calculated index values for sites as the

sum of the scores for their integrity components.

Majer and Beeston (1996) followed a similar

method to depict overall biological integrity of

ant assemblages in Australia.

The database created for each component above

displayed conditions for each 4th code HUC in

the basin assessment area. Components denoted

as number of species were then normalized in the

database to values ranging from 0 to 100 percent

by dividing the number of species in each HUC

by the total number of species possible in each

assessment component and multiplying by 100.

Components denoted as percentages were not

changed. In this way, all components were represented in the database as percentage values

within each 4th code HUC.

Then, for each of the integrity goals, all components were averaged by taking the arithmetic

mean of the percentages for each 4th code HUC.

The results, expressed as mean percentages in

each HUC, were mapped for each of the three

goals. These overall maps depict composite

conditions of all components under each goal.

The overall map for goal 1, maintaining species

viability, is shown in figure 62. Hydrologic unit

codes with higher percentage values in this map

represent locations with more threatened, endangered, endemic, rare, or candidate plants or animals, and overlap with the High Cascade Range

habitat corridor. Thus, the map highlights areas

that may warrant particular management attention

for this goal. Many HUCs ranked moderate (>20

percent mean condition) for goal 1. Several

Hydrologic unit codes ranked moderately high

(>40 percent), particularly in the Cascades Range

in central Washington, the Columbia River Gorge

and adjacent Washington-Oregon border, the Cascade Range in central Oregon, the Klamath Basin

in southern Oregon, the southern WashingtonIdaho border, and a portion of the northern Great

Basin in southeastern Oregon. These are areas

that may warrant particular management attention.

The overall map for goal 2, maintaining long-term

evolutionary potential of species, is shown in

figure 63. Hydrologic unit codes with higher percentage values in this map represent locations

with greater occurrence of disjunct species, and

with presence of hot spots and centers of concentration of biodiversity and species rarity or

endemism. As with the first goal, this map highlights areas that may warrant management attention. About half of the HUCs ranked at least

moderate (>20 percent mean condition) for goal

2. Several ranked moderately high (>40 percent),

particularly in the Cascade Range in southern

Washington, Glacier National Park and adjacent

lands in western Montana, the Idaho-Montana

border, the Blue Mountains and Hell’s Canyon

areas of the Washington-Oregon-Idaho conjunction, and parts of the Snake River Plain and

Owyhee Uplands in southern Idaho. There are

likely additional HUCs that rank moderately high

for goal 2 in southern Idaho for plant hot spots

that did not get identified.

The overall map for goal 3, managing for multiple

ecological domains and evolutionary timeframes,

is shown in figure 64. Hydrologic unit codes with

higher percentage values in this map represent

locations with greater occurrences of unique vertebrate assemblages, and locations with large carnivores (that is, the full trophic ladder). About

one-third of the HUCs ranked moderate (>20 percent mean condition) for goal 3, and none ranked

high. This is not surprising, as most of the components for this goal do not overlap, by definition of

unique geographic assemblages of species. Locations with moderately scored HUCs include

northwestern Montana, the Greater Yellowstone

105
Ecosystem in western Wyoming, the Cascades

Range in northcentral Washington, the Columbia

River Gorge, and much of the Klamath Basinnorthern Great Basin-Owyhee Uplands along

southern Oregon and Idaho. Collectively, these

areas cover some of the margins of the basin

assessment area where unique vertebrate assemblages would be expected to be found in such a

“land-locked” geography.

A cluster classification of HUCs (fig. 65) among

these three images revealed that goals 1 and 2 are

more similar than goal 3, as may be expected.

Maintaining species viability (goal 1) suggests

maintaining long-term evolutionary potential of

species (goal 2). All three goals and their associated HUC maps, however, provide a full, complementary set of conditions that may prove useful

for guiding ecosystem management planning at

the broad scale.

106
ICBEMP

Legend

Species = 0%

Species 1 − 20%

Species > 20 − 40%

Species > 40 − 60%

Species > 60 − 80%

Species > 80 − 100%

4th field hydrologic

units

State boundaries

Basin boundary

Figure 62—Composite map for ecological integrity goal 1, maintaining species viability, terrestrial components. Averagepercentage of all 4th-code hydrologic units in the basin assessment area among all components mapped for this goal (seefigs. 44-49).

107
ICBEMP

Legend

Species = 0%

Species 1 − 20%

Species > 20 − 40%

Species > 40 − 60%

Species > 60 − 80%

Species > 80 − 100%

4th field hydrologic

units

State boundaries

Basin boundary

Figure 63—Composite map for ecological integrity goal 2, maintaining long-term evolutionary potential of species, terrestrialcomponents. Average percentage of all 4th-code hydrologic units in the basin assessment area among all componentsmapped for this goal (see figs. 52 and 53).

108
ICBEMP

Legend

Species = 0%

Species 1 − 20%

Species > 20 − 40%

Species > 40 − 60%

Species > 60 − 80%

Species > 80 − 100%

4th field hydrologic

units

State boundaries

Basin boundary

Figure 64—Composite map for ecological integrity goal 3, managing for multiple ecological domains and evolutionarytimeframes, terrestrial components. Average percentage of all 4th-code hydrologic units in the basin assessment areaamong all components mapped for this goal (see figs. 54-61).

109
Figure 65—Cluster classification dendrogram of the three composite maps of ecological integrity goals (C1, C2,

and C3; in fig. 62, 63, and 64, respectively), among 4th-code hydrologic units within the basin assessment area.

Clustering was done by using complete linkage and farthest neighbor algorithms.

110
Conclusions and Discussion

Species Functions and Ecosystem

Processes

In this report, we have expanded on initial frameworks and findings (Marcot and others 1997) of

KEFs of species in the basin assessment area.

We tried to demonstrate basic ecological relations

among species and their environments that can

lead the way to greater confidence in managing

ecosystems for productivity and biodiversity. We

have not dealt with riparian, wetland, or aquatic

systems here, particularly with their unique constellations of species’ functions and ecosystem

processes. But they, too, are essential parts of

landscapes and need to be integrated with the

terrestrial portrait.

Our treatment of species functions—as with our

geographic scope—has been broad, and many

facets of ecological interactions of organisms

bear further telling and study. For example,

Willson (1996:101) lists six categories of direct

mutualisms (species mutually benefitting) of

organisms in terrestrial systems, including interactions among:

• Plants and animals (pollinators, dispersal

agents, woody plants and protective arthropods, and vascular plants with extrafloral

nectaries and animal protectors).

• Plants and fungi (mycorrhizal associations,

endophytic fungi that decrease attacks on

hosts by natural enemies, and lichens).

• Plants and micro-organisms (including

nitrogen-fixing microbes and angiosperms).

• Animals and other animals (plant-feeding

insects and ants, interspecific flocks and herds,

and other cases of animal-animal mutualisms).

• Animals and fungi (spore dispersal of

mycorrhizal fungi by small mammals, woodboring ambrosia beetles and fungal gardens).

• Animals and micro-organisms (digestive aids,

surface-dwelling microbes that provide protection with pathogens).

We have discussed some examples of many of

these mutualist species relations. Other commensal relations, as well as predatory, parasitic,

and other kinds of ecological relations, have

helped shape the composition and function of

ecological communities that we observe today.

The use of species function profiles and related

analyses of species’ KEFs can help to (1) determine the degree of functional diversity within

communities; (2) identify communities with high

(or low) redundancy in specific ecological functions, and therefore communities with high (or

low) resiliency or buffering capacity in the face of

disturbance regimes and systematic environmental

changes; and (3) identify ecological functions

with high (or low) redundancy among species, in

particular communities.1 Walker (1995) suggests

focusing conservation attention on functionally

important species groups with little redundancy;

that is, with few representative species. We may

extend his suggestion to include focusing conservation attention on communities with low redundancy in (or high variation among) ecological

functions. Risser (1995) notes that biodiversity

can affect ecosystem functions, and suggests

focusing initial management attention on the key

structuring processes at intermediate scales of

space and time.

1 All of these relations, particularly (b), need field

validation.

Chapter 5

111
Determining presence and redundancy of ecological functions also could be of help in posing specific goals for ecosystem restoration, particularly

for augmenting natural ecosystems with restored

ecosystems (Sinclair and others 1995) and by

matching functional diversity in ecosystems

undergoing restoration to that in undisturbed ecosystems. Examples of this include the use of species and ecosystem functions to monitor restoration progress in wetlands (Simenstad and Thom

1996) and to monitor trophic levels and feeding

functions (shredders, grazers, and predators) of

aquatic invertebrates in stream restoration projects (Murphy and Meehan 1991, Reeves and

others 1991). Among terrestrial communities of

the basin assessment area, studying patterns of

ecological functions (functional redundancy and

functional diversity) of forest seral stages targeted

for restoration, such as old single-stratum ponderosa pine forests, could prove useful as monitoring benchmarks or to help define management targets.

Bioindicators of Terrestrial

Ecosystem Health

In this report, we have further explored the potential use of lichens as bioindicators of various environmental situations, including old-growth forests, air quality, metal accumulation in soils, degree of soil acidity or alkalinity, fire and grazing

regimes, and other conditions. As we discussed

previously (Marcot and others 1997.), invertebrates also can serve as excellent early warning

signals of existing or impending changes in environmental conditions. Sampling a small part of

the invertebrate fauna, too, can help to index

diversity of the entire invertebrate fauna. For example, Oliver and Beattie (1996) discovered that

monitoring and assessment of terrestrial invertebrate biodiversity in Australian forests can be

done by careful use of invertebrate morphospecies

(species of the same morphologies)—principally

ants, beetles, and spiders. Their results also suggest which invertebrate taxa to use, as well as

sampling methods and sampling periods to yield

the most consistent and reliable monitoring. In

another study in the southern Appalachians,

Hollifield and Dimmick (1995) censused arthropod availability and abundance, and they inferred

from that suitability of logging roads, clearcuts,

and mature hardwood forests for ruffed grouse

(Bonasa umbellus). Other tests of plant or invertebrate indicators are presented by Nilsson and

others (1995), and Murtaugh (1996) discusses

statistical analyses of indicators. Although major

surveys and systematics studies are needed on

invertebrates in the basin assessment area, we feel

that such bioindicator approaches hold great

promise for rapid field evaluations of environmental conditions and of biodiversity in this

region as well.

Past and Future Changes

It is evident that the basin assessment area has

undergone tremendous prehistoric change in climate, habitats, and biota. Extinctions and originations of species, genera, families, and orders have

occurred over spans of millennia or longer. We

expect that natural extinctions, and hope that

originations (including speciations and natural

immigrations of native species), will proceed long

into the future. Less extreme than extinctions and

originations, we also expect changes in the geographic range of species’ abundance patterns over

time. For example, Kruckeberg and Rabinowitz

(1985) note that some rare plant taxa might become common, and some local endemics might

become widespread (although not as a rule), and

that the fossil record provides many instances of

widely distributed taxa that have become quite

constricted over time.

Patterns of evolution and extirpation of species

during the Quaternary cannot be discerned from

modern conditions. Conversely, recent historic

persistence of species probably cannot be used to

discern conditions over which species have persisted in the very long term or under which they

evolved. For example, some of the climatic regimes during the early Quaternary have no

modern analogs, and interglacial periods were

long and extreme enough to cause extinctions

but not long enough for significant plant evolution (Tausch and others 1993).

Pollen analysis from Wildcat Lake, Whitman

County, Washington suggests great changes in

aquatic and terrestrial environments over the past

1,000 years, with the most significant changes resulting from livestock introduction and subsequent alteration of natural vegetation, the

spread of weeds, and erosion, along with volcanism (Davis and others 1977). The response of

112
conifer vegetation to climate change may be rapid

in the more arid steppe communities of the basin

assessment area. Mack and Bryant (1974) report

much pine pollen in Quaternary steppe communities of the Columbia basin, thereby suggesting

potentially rapid dissemination and spread of pine

during periods of favorable climate.

More important and perhaps surprising, Fritts and

others (1979) report that climatic fluctuations in

the Columbia basin during the 17th through 19th

centuries differed significantly from those during

the 20th century and included large-scale variations. The range of recent historic climate does

not represent that of even just the past four centuries. Therefore, use of the range of natural historic variation in suggesting prehistoric climatic

conditions, and associated vegetation communities and conditions under which species persisted

or evolved, may need to be greatly revised.

We know that historic changes in vegetation conditions over the last two centuries have greatly

diminished habitats for some species and increased it for others. A combination of activities

(principally high-grade logging, forest fire suppression, and road building) have greatly enhanced habitat conditions for many species and

imperiled it for others. Lessons from this project

(chapters in Quigley and Arbelbide 1997) tell

us that recent land management changes and

growth of human populations in the basin assessment area have been accelerating, and that we

need to pay better attention to the broad-scale and

long-term additive effects of local extirpations

and local changes to habitats and environments.

Although global and regional climate changes

from increasing concentrations of greenhouse

gases in the lower atmosphere are beyond the

control of BLM and FS land planning, we can

nonetheless project potential outcomes and

inform publics and policymakers of effects.

One point of good news is that there is no evidence that acid precipitation is a major problem

for wildlife in the basin assessment area as it

is in other parts of the continent. Recent studies

by Vertucci and Corn (1996) suggest that highelevation aquatic amphibians—organisms highly

sensitive to and excellent indicators of acid precipitation and degradation of water quality—do

not seem to be incurring this problem in the

Rocky Mountains. But this does not mean that

populations of native amphibians are necessarily

stable; some indeed are declining for various other

reasons. Beyond the assessment area, amphibian

declines and disappearances have been linked to

quite a list of environmental woes including

chemical pollution, acid precipitation, increased

ultraviolet radiation, introduction of exotic species, pathogens, harvesting by humans, and natural population fluctuations (Blaustein and others

1994), as well as increasingly dessicating climates

(Pounds and Crump 1994) and depletion of

atmospheric ozone (Hileman 1994). We may

consider amphibians as crucial indicators of conditions to come and may use this list of environmental assailants to check our progress over time.

More localized current conditions and future

changes also can be heralded by heeding the

health of selected lichen, plant, and invertebrate

populations.

Ultimately, the productivity of habitats and their

biota depends on two main factors: solar energy

and soil. Solar insolation fuels photosynthesis

and primary productivity, which lie at the foundation of nearly all energy pyramids and food webs.

Virtually the only exceptions are the ancient

Archaebacteria—a group of methanogens, halogens, and extremely thermophilic acidophiles—

that stubbornly persist in high temperature and

pressure environments of thermal seeps in

Yellowstone, Mount St. Helens, and deep ocean

vents (black smokers)2 (Baross and Hoffman

1985). There is not much we can do about ensuring solar insolation for the rest of the terrestrial

biota, except to monitor changes in climate and

the atmosphere (for example, Beerling and others

1995, Dubayah 1994), perhaps by using lichens

and bryophytes as litmus tests of air quality.

The soil, however, is literally at our feet and

figuratively at the hand of management, and there

is much we can do to monitor and guide its health.

It is ironic that soil health is so influenced by

microbes but manifests its effect over vast geographic areas. Its complement of macro- and

micro-organisms and their vast array of ecological

functions can be quickly depleted under careless

management yet may take so much longer to rejuvenate. And results of site-specific management

2 Personal communication. 1996. P. Frenzen, Mount St.

Helens National Volcanic Monument, USDA Forest

Service, Washington.

113
activities that influence soil compaction, organic

matter input, dispersal of mycorrhizae, vitality of

seed banks, and persistence of microbiotic crusts,

can be monitored with satellites. Maintaining

ecological integrity will likely include heeding the

lessons of soil by maintaining rich belowground

biota with their full complement of ecological

functions. In soil lies the vitality of future resources and ecosystems.

Managing for Ecological Integrity

We have identified locations within the basin assessment area that may invite attention of policy

planners for meeting the three ecological goals

for long-term integrity of terrestrial ecosystems

at the broad scale. Terrestrial ecosystem integrity

would likely benefit from further, similar assessments of our integrity components at finer resolutions for use in mid-scale planning and management activities.

Taken with assessments presented in Marcot and

others (1997) and analyses in this report, we are

encouraged that good opportunities still exist in

the basin assessment area for meeting all three

goals of terrestrial ecological integrity (that is,

to maintain species viability, maintain long-term

evolutionary potential of species, and manage for

multiple ecological domains and evolutionary

timeframes). To meet these goals, however, careful attention needs to be paid in the short term to

those integrity components most at risk. These

include threatened, endangered, rare, and potentially rare plants and animals; providing for species’ habitats at the peripheries of their ranges

and in disjunct population locations; providing

for conditions to maintain hot spots and centers

of concentration of plant and animal biodiversity

and species rarity and endemism; and providing

for habitats to maintain unique assemblages of

plants and animals.

Other ecosystem integrity components of potential interest to successful ecosystem management

include ensuring long-term productivity of soils

and the belowground world; and attending to

scarce and still-diminishing vegetation conditions,

principally rare plant communities, native grasslands, native shrublands, and late seral subalpine

multilayer forests. We expect, with careful salvage thinning and logging of many lower elevation forests, along with prudent reintroduction of

fire into these ecosystems, that late seral ponderosa pine single-layer forests eventually can be encouraged to return. Additional components of

ecological integrity and terrestrial species conservation that may be of interest to managers in the

basin assessment area are presented by Marcot

and others (1997).

An Uncertain Future for Humans

and Terrestrial Ecosystems: Making

Tough but Informed Choices

“The extirpation of populations and species of

organisms exerts its primary impact on society

through the impairment of ecosystem services”

(Ehrlich 1988:24). Terrestrial ecosystems continue to surprise us. Like the intricate layering of

Hindu temple architecture, on closer inspection

they reveal new details and structures and new

services to aid human societies. For example,

pharmacological and economic benefits of maintaining the full expression of plant diversity have

hardly been explored in terrestrial ecosystems of

the basin assessment area, but other experiences

(for example, Aylward and Fendt 1995) suggest a

richness yet to be realized. Likewise, we have

only begun to recognize the long-term value to

human societies—both native and “naturalized”—

in maintaining the full diversity and productivity

of terrestrial systems, principally in soils and the

belowground world. Again, experiences elsewhere (for example, papers in Bissonette and

Krausman 1995) shed great light on how sustainable use of natural resources can be provided by

better understanding ecosystem processes and

living within those means—or suffering the consequences of unnecessary resource scarcity, economic disruption, and social uncertainty (Rathore

1995).

It is not entirely clear what future effects will

come of increasing human occupation of the basin

assessment area, particularly from increases in

roads, recreation, and human habitations in the

“urban-forest interface.” Although, on average,

human population density of the basin assessment area is low compared with other parts of

the United States (McCool and others 1997),

much of the assessment area is already roaded

and under active land management for various

goods and services (fig. 3). Rather certain scenarios of increasing human populations (McCool and

114
others 1997) and rather uncertain scenarios of

regional climate change together could pose

grave problems and challenges to managers for

maintaining a strong semblance of native biota,

ecological processes, and ecosystems. Assuredly,

terrestrial ecosystems of the basin assessment

area will continue to exist in one form or another.

But, it is not clear how ecosystem processes such

as nutrient cycling and frequency of wildfires,

structures such as plant and animal communities

and soil microbiota, and services such as soil productivity, nutrient storage, and interception of

surface water runoff will be altered by increasing

human presence and changing Federal land management policies and activities.

Human occupation of the land often serves to

sharpen boundaries between managed and unmanaged landscapes. Examples of such boundaries are rural human habitations adjacent to

National Forest lands; silviculturally managed

timber stands adjacent to old-growth forests or research natural areas; and grazing pastureland or

agricultural fields adjacent to native Agropyron

or Fescue grasslands or sagebrush steppe. Managed systems are characterized by very large

fluxes in exports of soil, nutrients, pesticides,

and inorganic ions, along gradients steeper than

naturally occur. These changes in rates ultimately

affect soils, biota, and atmospheric exchange

rates (Correll 1991). In soils, pH, Eh, ionic composition, nutrient and toxic metal concentrations,

and organic matter pathways can be altered

(Correll 1991). Such are the kinds of complex

ecosystem changes that intensive natural resource management may induce and that managers may wish to better quantify.

Our aim in this report was to contribute to understanding the history and ecology of species, communities, and ecosystems of the basin assessment

area. We hope we have helped build a basis for

asking the right kinds of ecological questions, and

a framework for beginning to answer those questions when conducting ecosystem planning and

management. We also hope that the era of terrestrial ecosystem management on Federal public

lands has evolved far beyond a simple focus on

threatened and endangered species and big game,

although these too are legitimate and essential

parts of ecosystem planning. Many ecological

relations among species affect diversity, productivity, and sustainability of ecosystems. Some

relations are subject to change and can be

threatened just as individual species or populations can, and thereby also warrant exquisite

attention in research and management. We hope

that the fruitless arguments of “species versus

ecosystems” have been finally interred—as both

are obviously essential and inextricable foci for

understanding and for successfully managing

ecosystems.

As well, an understanding of past and current

situations—including variations in taxonomic

groups ecological functions of species, redundancy of functions among species and communities, turnover (loss and gain) of species and taxa,

and changes in climates and disturbance regimes—is fundamental to estimating the natural

range of conditions, historic and present. Kay

(1994) concludes that, historically, Native Americans commonly determined the structure of entire

plant and animal communities by hunting and by

setting fires (also see papers cited in Knowles and

Knowles 1993), and that a current “natural regulation” approach to management does not recognize

and thus probably would not replicate such historic conditions. He also pointed out that the historic reductions in number and distribution of

beaver (Castor canadensis) throughout the West

has had drastic effects on declining distribution of

some wetland and upland ecological communities.

Along with others, Kay asks the question, “What

is natural?” It is a question we must answer within

contexts of ecosystem policy goals, historic conditions, an understanding of prehistoric changes,

background rates of extinction and origination of

organisms at several taxonomic levels, the current

biophysical template of the land, and even societal

and political desires.

Ultimately, the range of natural conditions and

ecosystem dynamics—if defined through a standard, rigorous, and repeatable process—spells

out the history of the land and might provide an

insight into what the land is capable of supporting

in the future. Our choice as society is then to adhere to the dimensions of that envelope, or to

force ecosystems into new, and usually more reduced, constitutions. At least, we hope, for terrestrial ecosystems of the basin assessment area,

it will be a learned and informed decision.

115
Acknowledgments

Tom Quigley and Gary Benson provided policy

reviews of the manuscript. Becky Gravenmier

and her Spatial Staff of the Science Integration

Team provided estimates of the size of the assessment area and vegetation communities. Cary

Lorimor produced the maps. Ted Fremd, chief

paleontologist, John Day Fossil Beds National

Monument, graciously provided species lists from

his ongoing studies, from which we developed

our analyses and discussions on paleoecological

trends of vertebrate communities. Jonathan

Marcot provided additional field paleoecological

interpretation. Jeff Jones provided analyses of

rates of historic change of vegetation communities and helped with interpretation of these data.

Sue Ferguson provided data on climate change

model results and helped with interpretations of

these data.

English Equivalents

1 kilometer (km) = 0.62 mile

1 meter (m) = 3.281 feet

1 hectare (ha) = 2.47 acres

Degrees Celsius (o

C) = 5/9(oF-32),

where o

F = degrees Fahrenheit

116
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Biodiversity in managed landscapes theory and practice. New York: Oxford University Press:

96-107.

Wilson, D.E.; Reeder, D.M., eds. 1993. Mammal species of the world: a taxonomic and geographic

reference. 2d ed. Washington, DC: Smithsonian Institution Press.

Wilson, E.O. 1988. The current state of biological diversity. In: Wilson, E.O.; Peter, F.M., eds.

Biodiversity. Washington, DC: National Academy Press: 21-27.

Windsor, D.A. 1995/96. Endangered interrelationships: the ecological cost of parasites lost. Wild

Earth. 5(4): 78-83.

125
Appendix 1

Categories of abundance and breeding or migration status of birds in the

basin assessment area, as denoted in the species-environment relations

database.

Abundance of birds:

A = Abundant

Within the right habitat and season, birds in this category would seem to be ubiquitous and numerous.

C = Common

Within the right habitat and season, birds in this category would be present.

U = Uncommon

Unlikely that birds in this category would be seen unless specifically searched for in the correct

location and time. Populations are larger than those of rare birds but still not always seen when

expected.

R = Rare

Seen every year but in very low numbers, usually less than 10 sightings per year. The Pacific loon, for

example, is rare in Washington. Whereas the Pacific loon is out of its normal range when sighted

inland, other species can be rare within their breeding range. For example, the black-backed

woodpecker is a year-round resident of Montana. It breeds there but is at the edge of its range and is

rarely seen because its populations are small.

I = Irregular

Usually seen in low numbers and not seen every year. An example is the Pacific loon along the

Columbia River in Oregon; this species is seen in this location only in some years and always in low

numbers. Another example is snowy owl, which is uncommon in the inland West when winters are

harsh farther north, but which is absent in other years.

Breeding status of birds:

R = Resident

A bird species that breeds in the state and can be seen there year-round.

B = Breeding

A species that breeds in the state but winters elsewhere.

N = Nonbreeding summer resident

An unusual situation. Only a few bird species in the inland West fall into this category. One example

is the blue jay, which does not breed anywhere in the inland West, but quite a few occur all summer in

northwestern Wyoming.

W = Wintering

Bird species that spend the winter and breed elsewhere.

M = Migrants

Bird species present only in the spring and fall as they move between wintering and breeding areas.

126
Appendix 2

Lichen and bryophte species groups as bioindicators. Additional comments

included here on species groups are from the Plant Task Group, SIT,

Interior Columbia Basin Ecosystem Management Project (see Marcot and

others 1997 for listings of specific species in each group).

Air Quality Indicators

Lichen Groups:

Bog lichens

Calcareous indicator lichens (a large group of species)

Calcareous steppe indicator lichens

Charred snag lichens (indicates recent fire)

Excess nitrogen indicator lichens

Fencepost lichens

Forage lichens

Fruticose tree lichens (only some of these species are old-growth indicators)

Leaf lichens (this is a large group of variable species)

Metal-rich indicator lichens

Moss and ditritus binders lichens

Nitrogen-fixing epiphytes lichens

Nitrogen-fixing rock lichens

Nitrogen-fixing soil lichens

Oceanic forage lichens

Oceanic leaf lichens

Oceanic log lichens

Oceanic rock lichens

Oceanic tree crust lichens

Pin lichens

Pioneer soil stabilizers lichens

Riparian lichens (indicators of high humidity)

Rock crusts lichens

Rock macro lichens (many species in this group)

Rotten log and tree base lichens (indicates ecological continuity)

Seepage lichens

Sheltered ledges and overhangs lichens

Soil lichens

Steppe soil crust lichens

Tree crusts lichens

Tundra forage lichens

Tundra rock lichens

Tundra sod builders (some species are calcareous indicators)

Urban pollution-tolerant lichens

Vagrant ground lichens

Bryophyte Groups:

Epiphytic bryophyte

Humus duff bryophyte

Peatlands bryophtye

Rock other bryophyte

127
Rock wet bryophyte

Soil alkaline bryophyte

Soil dry bryophyte

Soil wet bryophyte

Alkaline (Carbonate) soil indicators

Lichen Groups:

Calcareous indicator lichens (large groups of species)

Calcareous steppe indicator lichens

Rock crusts lichens

Tundra sod builders (some species are calcareous indicators)

Vagrant ground lichens

Carex calcareous peatlands group (perennially saturated peat; most areas are open, but sedges on the

edges of these peatlands are in partial shade)

Bryophyte Groups:

Aquatic submerged bryophyte

Peatlands bryophtye

Rock calcareous bryophyte

Rock wet bryophyte

Soil dry bryophyte

Soil wet bryophyte

High waterflow indicators

Lichen Groups:

Aquatic lichens (good indicators of water quality and flow levels)

Seepage lichens

Bryophyte Groups:

Aquatic submerged bryophyte

Rock wet bryophyte

Soil alkaline bryophyte

Low waterflow indicators

Lichen Groups:

Aquatic lichens (good indicators of water quality and flow levels)

Seepage lichens

Bryophyte Groups:

Aquatic submerged bryophyte

Rock wet bryophyte

Soil alkaline bryophyte

Grazing indicators-decreasers

Lichen Groups:

Calcareous steppe indicator lichens

Nitrogen-fixing soil lichens

Soil lichens

Steppe soil crust lichens

128
Tundra forage lichens

Tundra sod builders (some species are calcareous indicators)

Vagrant ground lichens

Carex sagebrush steppe dry group

Carex sagebrush steppe wetland riparian group (hot dry summers)

Grazing indicators-increasers

Lichen Group:

Pioneer soil stabilizers lichens

Bryophyte Groups:

Aquatic submerged bryophyte

Peatlands bryophtye

Rock wet bryophyte

Soil alkaline bryophyte

Soil dry bryophyte

Soil wet bryophyte

High-nitrogen indicators

Lichen Groups:

Excess nitrogen indicator lichens

Fencepost lichens

Rock crusts lichens

Rock macrolichens (many species in this group)

Urban pollution-tolerant lichens

Bryophyte Groups:

Aquatic submerged bryophyte

Rock other bryophyte

Rock wet bryophyte

Soil alkaline bryophyte

Soil dry bryophyte

Soil wet bryophyte

Metal-rich rock indicators

Lichen Groups:

Metal-rich indicator lichens

Rock crusts lichens

Bryophyte Groups:

Aquatic submerged bryophyte

Rock other bryophyte

Rock wet bryophyte

Soil alkaline bryophyte

Soil dry bryophyte

Soil wet bryophyte

129
Old-growth forest indicators

Lichen Groups:

Calcareous steppe indicator lichens

Charred snag lichens (indicates recent fire)

Forage lichens

Fruticose tree lichens (only some of these species are old-growth indicators)

Moss and ditritus binders lichens

Nitrogen-fixing epiphytes lichens

Oceanic forage lichens

Oceanic leaf lichens

Oceanic log lichens

Oceanic tree crust lichens

Pin lichens

Riparian lichens (indicators of high humidity)

Rotten log and tree base lichens (indicates ecological continuity)

Steppe soil crust lichens

Tree crusts lichens

Bryophyte Groups:

Decayed wood bryophyte

Epiphytic bryophyte

Humus duff bryophyte

Peatlands bryophtye

Rock calcareous bryophyte

Rock other bryophyte

Rock wet bryophyte

Soil dry bryophyte

Soil wet bryophyte

High soil temperature indicators

Bryophyte Groups:

Rock calcareous bryophyte

Rock other bryophyte

Rock wet bryophyte

Soil alkaline bryophyte

Soil dry bryophyte

Soil wet bryophyte

Low soil temperature indicators

Bryophyte Groups:

Peatlands bryophtye

Rock calcareous bryophyte

Rock other bryophyte

Rock wet bryophyte

Soil dry bryophyte

Soil wet bryophyte

Tundra forage lichens

Tundra sod builders (some species are calcareous indicators)

Vagrant ground lichens

130
Carex calcareous peatlands group (perennially saturated peat; most areas are open but sedges on the

edges of these peatlands are in partial shade)

Carex noncalcareous peatlands group (perennially saturated peat; most areas are open but sedges on

the edges of these peatlands are in partial shade)

Coarse soil texture indicators

Bryophyte Groups:

Peatlands bryophtye

Rock calcareous bryophyte

Rock other bryophyte

Rock wet bryophyte

Soil alkaline bryophyte

Soil dry bryophyte

Soil wet bryophyte

Vascular Plant Groups:

Penstemon acuminatus group (sandy dunes and sandy areas near rivers)

Penstemon foothills to montane dry rocky group (rocky margins of old stream channels, scree, rock

walls, outcrops, rocky roadcuts, and well drained areas; indicators of dry conditions)

Penstemon foothills to montane meadow group (indicators of changes in hydrologic regime)

Fine Soil Texture Indicators

Lichen Group:

Steppe soil crust lichens

Bryophyte Groups:

Peatlands bryophtye

Rock calcareous bryophyte

Soil alkaline bryophyte

Soil dry bryophyte

Soil wet bryophyte

131
Appendix 3

Rare or potentially rare vascular plant species as bioindicators. Note that

this list includes only the rare or potentially rare species explicitly

considered in this assessment. Other indicator plants that are not rare or

potentially rare also can be identified. Additional comments included here

on species are from the Plant Task Group, Science Integration Team,

Interior Columbia Basin Ecosystem Management Project.

Alkaline (Carbonate) Soil Indicators:

Arabis fecunda

Astragalus paysonii

Botrychium ascendens

Botrychium lineare

Botrychium pedunculosum

Carex lenticularis var. dolia

Carex parryana ssp. idahoa

Chrysothamnus parryi ssp. montanus

Erigeron lackschewitzii

Lesquerella carinata var. languida

Lesquerella paysonii

Penstemon compactus

Physaria integrifolia var. monticola

High WaterFlow Indicators:

Artemisia ludoviciana ssp. estesii

Gratiola heterosepala

Grindelia howellii

Oxytropis campestris var. columbiana

Grazing Indicators - Decreasers (sensitive to grazing disturbances):

Astragalus sinuatus

Botrychium ascendens

Botrychium crenulatum

Carex parryana ssp. idahoa

Cymopterus acaulis var. greeleyorum

Eriogonum novonudum

Gratiola heterosepala

Lesquerella carinata var. languida

Lesquerella paysonii

Mimulus evanescens

Mimulus patulus

Oryzopsis contracta

Phacelia lutea var. calva

Phacelia lutea var. mackenzeorum

Silene spaldingii

Stanleya confertiflora

Thelypodium howellii ssp. howellii

Thelypodium howellii ssp. spectabilis

Metal-Rich Rock Indicators:

132
Haplopappus radiatus

Penstemon nikei

High Soil Temperature Indicators:

Astragalus diaphanus var. diurnis

Lesquerella carinata var. languida

Oryzopsis contracta

Low Soil Temperature Indicators:

Carex lenticularis var. dolia

Castilleja pilosa var. steenensis

Lesquerella humilis

Coarse Soil Texture Indicators:

Allium aaseae

Allium constrictum

Allium punctum

Allium robinsonii

Astragalus columbianus

Astragalus diaphanus var. diaphanus

Astragalus diaphanus var. diurnis

Astragalus mulfordiae

Astragalus pulsiferae var. suksdorfii

Astragalus sinuatus

Astragalus yoder-williamsii

Botrychium lunaria

Carex lenticularis var. dolia

Chrysothamnus parryi ssp. montanus

Cypripedium fasciculatum

Descurainia torulosa

Erigeron lackschewitzii

Hackelia cronquistii

Hackelia venusta

Lesquerella carinata var. languida

Lesquerella humilis

Lesquerella paysonii

Oxytropis campestris var. columbiana

Papaver pygmaeum

Penstemon glaucinus

Penstemon kingii

Penstemon lemhiensis (does well in disturbed subsoils, but seems to do best on developed, dark soils

[mollisols] of gravelly loam texture)

Silene spaldingii

Stanleya confertiflora

Fine Soil Texture Indicators:

Aster jessicae

Astragalus sinuatus

Botrychium ascendens

Botrychium crenulatum

Botrychium lunaria

Botrychium paradoxum

133
Botrychium pedunculosum (successional relations unclear; species may prefer late seral grassland

habitats, as well as early to mid successional forest habitat)

Calochortus longebarbatus var. longebarbatus (soils are colluvial)

Calochortus longebarbatus var. peckii

Calochortus nitidus

Camissonia pygmaea (plants at soil interface adjacent to talus)

Carex parryana ssp. idahoa (occurs in minerotrophic [rich] subalpine fens consisting of sedge peat;

persistent, saturated emergent wetlands)

Chaenactis cusickii (ash deposits weathering to clays; few associates, almost always annuals; dry,

barren sites)

Erigeron lackschewitzii (found exclusively on calcareous soil derived from dolomite; soils are

cryptorthents of gravelly silt or loam)

Eriogonum novonudum

Grindelia howellii (plant occurs in many disturbed sites, but its native habitat may be upper drawdown

zones of prairie ponds in the Ovando Valley)

Haplopappus liatriformis (does poorly in areas of significant grazing)

Howellia aquatilis (ponds surrounded by narrow ring of cottonwood and tall deciduous shrubs;

populations fluctuate widely among years because of variation in precipitation; recent wild

fluctuations of precipitation [1988-drought; 1993-wet; 1994-drought] have reduced populations)

Lepidium davisii (desert playas; hard gray-white clays; pH 6-7 at all playas; very few associated

species; barren; roots deep in cracks)

Primula alcalina

Rorippa columbiae

Silene spaldingii (biscuits in biscuit-scabland on loess, alluvium, aeolian soils)

Trifolium douglasii (little known about this species)

134
Index

A

acid precipitation 113

air quality 35

alpine communities 54

alpine tundra 58, 60

Archaebacteria 113

B

Basin and Range 20

basin assessment area 2

beta diversity 65

big game 115

biodiversity 18

bioindicators 33, 112

biophysical factors 28

birds 42

bivalves 35

Blue Mountains 20

broadscale (defined) 1

brood parasitism 44

bryophyte groups 23

C

candidate species 74, 76

carnivores 96

carrion feeders 52

Cascade Range 2, 20

centers of concentration of biodiversity 90

Clearwater Maritime refugium 27

climate change 27, 57

CO2 63

Columbia Plateau 20

connectivity of habitats 76

cowbirds 44

D

disease 39

disharmonious faunas 65

disjunct populations 27, 87

dispersal 36

dissemination 28

E

ecological domains90

ecological integrity74, 114

ecological reporting units3, 96

ecosystem health112

ecosystem management 1

ecotones 63

endemic plants 25

endemic vertebrates 76

evolution 112

evolutionary timeframe 90

extinction 65, 112

extirpation 112

F

fire regimes 60

fire suppression 11

Forest Ecosystem Management

Assessment Team [FEMAT] 2

fossil record 70

functional diversity 16, 112

functional redundancy 49, 55, 56

functionally analogous species 15

G

gastropods 35

glacial cycles 65

Great Basin 20, 62, 64

H

habitat corridors 87

herbivory 52

Holocene Recent epochs 64

hot spots 59, 90

human population density 114

hydrologic unit codes (HUCs) (defined) 1

I

indicators 33

invertebrates 35

J

John Day Fossil Beds National Monument 66

K

key ecological functions 49

key environmental correlates 14

keystone mutualists 33

L

landform 3

lichen groups 23

life forms 20

locally endemic vertebrates 76

135
M

macroecology (defined) 1

megafaunal extinction 65

micro-organisms 35, 63

microbiotic crusts 114

microfungi 20

midscale (defined) 1

Mojave 59

mollusks 35

multiple ecological domains 90

mushrooms 20

mycorrhizae 20, 60, 63, 114

N

Native Americans 115

natural 115

neoendemics 26

Neotropical migratory songbirds 44

nitrogen cycles 39, 60

nonanalogue communities 65

Northwest Forest Plan 2

nutrient cycling relations 53

O

Okanogan Highlands 20

old-growth forest 35

overgrazing 35

ownership 3

P

paleoecology 64

paleoendemics 26

Palouse 25

parasites 39

phoresis 36

phoretic dispersal 36

plant dispersal modes 28

plant pollinators 33

plants and allies 20

pleistocene 26, 64, 65

pollination ecology 33

pollination vectors 53

polypores 20

primary burrow excavators 53

primary cavity excavators 53

puffballs 20

Q

Quaternary period 64, 112

R

range of historic conditions 65

range of natural historic variation 113

rare plants 76

redundancy of ecological functions 112

regional climate change 115

resupinate 20

S

semiendemic species 42

slugs 35

snails 35

soil conditions 35

soil health 113

soil temperature 35

soil textures 35

soils 23

species function profiles 15, 44, 49, 52, 76, 111

species groups 14

species-environment relations

(SER) database 14

subalpine forest communities 58, 60

subbasins (defined) 3

subwatersheds (defined) 3

T

temporal continuity 35

terrestrial species (defined) 1

Tertiary period 66

threatened and endangered plants 76

threatened and endangered species 115

threatened and endangered

vertebrate species 75

total functional diversity (also

see functional diversity) 16, 49, 56

truffles 20

V

vascular plant groups 23

vascular plant species 23

vegetation classifications 3

vegetation communities 3

vegetation cover types 3

vertebrate species endemism 40

vertebrates 40

viability 33, 74

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Marcot, B.G.; Croft, L.K.; Lehmkuhl, J.F.; Naney, R.H.; Niwa, C.G.; Owen, W.R.;

Sandquist, R.E. 1998. Macroecology, paleoecology, and ecological integrity of

terrestrial species and communities of the interior Columbia River basin and northern

portions of the Klamath and Great Basins. Gen. Tech. Rep. PNW-GTR-410. Portland,

OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research

Station. 136 p.

This report presents information on biogeography and broad-scale ecology (macroecology)

of selected fungi, lichens, bryophytes, vascular plants, invertebrates, and vertebrates of the

interior Columbia River basin and adjacent areas. Rare plants include many endemics

associated with local conditions. Potential plant and invertebrate bioindicators are identified.

Species ecological functions differ among communities and variously affect ecosystem

diversity and productivity. Species of alpine and subalpine communities are identified that

may be at risk from climate change. Maps of terrestrial ecological integrity are presented.

Keywords: Macroecology, paleoecology, ecological integrity, terrestrial communities,

ecosystems, wildlife, fungi, lichens, bryophytes, vascular plants, invertebrates, arthropods,

mollusks, amphibians, reptiles, birds, mammals, endemism, interior Columbia River basin,

Klamath Basin, Great Basin.

The Forest Service of the U.S. Department of

Agriculture is dedicated to the principle of multiple

use management of the Nation’s forest resources

for sustained yields of wood, water, forage, wildlife,

and recreation. Through forestry research, cooperation with the States and private forest owners, and

management of the National Forests and National

Grasslands, it strives—as directed by Congress—to

provide increasingly greater service to a growing Nation.

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etc.) should contact the USDA’s TARGET Center at

(202) 720-2600 (voice and TDD).

To file a complaint, write the Secretary of Agriculture,

U.S. Department of Agriculture, Washington, DC 20250,

or call (800) 245-6340 (voice), or (202) 720-1127 (TDD).

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