An Annotated Bibliography of 

Scientifi c Literature on 

Managing Forests for 

Carbon Benefi ts

Sarah J. Hines

Linda S. Heath

Richard A. Birdsey

United States

Department of 

Agriculture

Forest Service

Northern 

Research Station

General Technical

Report NRS-57
Abstract

Managing forests for carbon benefi ts is a consideration for climate change, bioenergy, 

sustainability, and ecosystem services. A rapidly growing body of scientifi c literature on forest 

carbon management includes experimental, modeling, and synthesis approaches, at the standto landscape- to continental-level. We conducted a search of the scientifi c literature on the topic 

of managing forests for carbon, and compiled an annotated list of citations. We chose to focus 

specifi cally on studies that addressed carbon in aboveground carbon pools, at both the micro (tree, 

stand) and macro (landscape, policy) levels. Aboveground pools include: live tree, understory, 

standing dead wood, down dead wood, and forest fl oor. The temporal scope of the literature search 

was the period 2000-2008 and the geographical scope was the temperate and boreal forests 

mainly in the United States, but also Canada, Europe, Russia, Japan, China, New Zealand, and 

Australia. The annotated bibliography is available at: http://www.nrs.fs.fed.us/pubs/ as a printable 

document; the references included in the bibliography are also available as a downloadable 

EndnoteTM database or basic text fi le. The CD-ROM included with this publication contains the 

datasase and text fi les.

Visit our homepage at: http://www.nrs.fs.fed.us/

Published by: For additional copies:

U.S. FOREST SERVICE U.S. Forest Service 

11 CAMPUS BLVD SUITE 200 Publications Distribution

NEWTOWN SQUARE PA 19073-3294 359 Main Road

 Delaware, OH 43015-8640

February 2010 Fax: (740)368-0152

The Authors

SARAH J. HINES (shines@fs.fed.us) is an ecosystem services specialist with the U.S. Forest 

Service Northern Research Station (NRS) and Northeastern Area State & Private Forestry (NA), 

both located at Newtown Square, PA. LINDA S. HEATH (lheath@fs.fed.us) is research forester, 

with NRS at Durham, NH. RICHARD A. BIRDSEY is a program manager with NRS at Newtown 

Square, PA.

Illustration on the cover is by Eric Fiegenbaum

The use of trade, fi rm or computer names in this publication is for the information and convenience 

of the reader. Such use does not constitute an offi cial endorsement by the USDA Forest Service of 

any product or service to the exclusion of others that may be suitable.

Manuscript received for publication 30 June 2009
1

INTRODUCTION

Scientists and many policy-makers acknowledge that 

land-use change and forestry play a signifi cant role in 

the global carbon cycle. Deforestation, which is a type 

of land-use change, and poor management practices 

such as degradation of global forests contributed 17.4 

percent of global greenhouse gas releases in 2004 

(IPCC 2007). When sustainably managed, forests 

can help mitigate climate change by sequestering 

atmospheric carbon dioxide in biomass and soils. In 

the United States, forests and products are estimated 

to sequester a net 745 Tg CO2 under current land 

use and management practices (U.S. EPA 2008), 

off setting as much as 20 percent of U.S. fossil fuel 

emissions (U.S. CCSP 2008a). Estimates suggest that 

the potential sequestration for U.S. forests may be in 

the range of 1500 to 2300 Tg CO2 per year (Heath et 

al. 2003).

Th is bibliography represents a compilation of literature 

on the topic of the eff ects of forest management on 

carbon stocks or fl uxes published in the years 2000-

2008. It is intended as a starting point for scientists 

and managers regarding forest-based mitigation of 

climate change, as well as background information 

at the strategic level for mitigation opportunities. 

Pronounced climate change eff ects on our nation’s 

forests and grasslands are beginning to be noticed 

(U.S. CCSP 2008b), so land managers are under 

increasing pressure and scrutiny to manage forests 

to be resilient to, and preventive of, climate change. 

What management practices, employed in which 

scenarios, can contribute to mitigation? Th is question 

is complex with informed answers often requiring a 

combination of global, regional, and local-scale data, 

and an intimate understanding of forest management 

practices and options considered with the uncertainty 

of climate change and other natural disturbances. 

Managers of public lands with multiple-use mandates 

are required to balance and prioritize multiple, 

sometimes competing objectives. In some regions or 

for some ownerships, mitigation practices may focus 

on maximizing landscape-level ecosystem carbon 

stocks while considering uncontrollable disturbances, 

while in other regions or for other landowners, 

objectives may focus on sustaining or increasing 

carbon sequestration in wood and bioenergy products 

which are a substitution for fossil fuel intensive 

products. In all scenarios, land managers may question 

whether, when, and how management practices such 

as thinning and fuel treatments create net benefi ts for 

the climate, or whether amendments should be used 

to enhance growth and productivity. In Figure 1, the 

red arrows depict emissions and sequestration by or 

within the forestry sector; the green arrows represent 

emissions outside of this sector. It should be noted 

that biofuels that substitute for fossil fuels prevent the 

release of fossil fuel-related emissions.

Figure 1.—Forest sector 

mitigation strategies 

need to be assessed with 

regard to their impacts 

on carbon storage in 

forest ecosystems, on 

sustainable harvest 

rates, and on net GHG 

emissions across all 

sectors. (Figure from 

Nabuurs et al. 2007, 

used with permission.)
2

THE BIBLIOGRAPHY

Purpose

Th is annotated bibliography is intended primarily 

as a reference for land managers who are seeking to 

understand the eff ects of forest management activities 

and climate-induced changes on the carbon stocks and 

fl uxes of a stand or landscape. Each study listed in the 

bibliography is followed by a concise summary of its 

methods and primary results. Th e compilation should 

be helpful to managers of federal land and wildlife 

management agencies, and managers in the private 

sector, to show their land management decisions are 

taking into account ‘best available science’ (APA 1946, 

U.S. GAO 2003, NFMA 1976). At a strategic level, 

the Resources Planning Act (1974) requires the U.S. 

Secretary of Agriculture to conduct an assessment of the 

Nation’s renewable resources every 10 years. Subsequent 

amendments require an analysis of the rural and urban 

forestry opportunities to mitigate the buildup of 

atmospheric carbon dioxide and reduce the risk of global 

climate change. Th is compilation helps identify forest 

management opportunities for the RPA Assessment.

Scope

Th is bibliography includes studies and papers which 

examine the eff ects of forest management activities on 

all aboveground forest carbon pools, at both the micro 

(tree, stand) and macro (landscape, policy) levels. 

Aboveground pools include: live tree, understory, 

standing dead wood, down dead wood, and forest 

fl oor. Aboveground pools are generally more dynamic 

than the soil carbon pool and are more dramatically 

aff ected by near-term management activities. Soil 

carbon is a critically important component of the 

terrestrial carbon pool and can be dramatically altered 

by land-use change, but soil carbon studies are often 

specialized and focus only on soil carbon, whereas 

our focus is on the entire forest. Th ere are existing 

bibliographies of forest management and its eff ects 

on soil carbon (Heath and Smith 2000, Johnson and 

Curtis 2001, Hoover 2003), as well as an extensive 

discussion of the potential of U.S. forests to sequester 

carbon (Kimble et al. 2003), so our focus is on 

studies that include all forest pools or at least most 

aboveground carbon pools.

Because continued forest-based mitigation of 

climate change is dependent upon forests’ successful 

adaptation to climate change, the literature and 

consequently this bibliography also includes limited 

related studies on forest management for adaptation. 

Th e geographic scope includes temperate and 

boreal forests in North America, Europe, Russia, 

Japan, China, New Zealand, and Australia. Forest 

management activities included in this bibliography 

consist of: aff orestation/reforestation, amendment 

application, avoided deforestation, fuels treatment, 

harvesting, rotation length, general management 

intensity decisions, and species selection decisions. 

Incidental or involuntary climate eff ects include 

climate-driven natural disturbances (fi re, insects or 

disease), involuntary fertilization (carbon dioxide and 

nitrogen deposition), changing weather patterns, and 

changing climate (temperature and precipitation) 

over the long term. Publications are also drawn from 

optimization studies; carbon fl ux analyses; studies of 

aff orestation, reforestation, or avoided deforestation; 

economic analyses; and climate change feedback and 

eff ects.

At the time of publication, this bibliography contains 

171 publications sorted into fi ve broad categories: 

1) experiments and observations, 2) models, 3) 

experiments with some pools modeled, 4) literature 

reviews and meta-analyses, and 5) policy perspectives. 

All publications listed include brief summaries of 

fi ndings relevant to managing forests for carbon, 

especially aimed at forest-based mitigation of climate 

change. To the extent that they provide carbon pool 

data, carbon fl ux studies have been included (e.g., 

Pacala et al. 2001). And to the extent that they focus 

substantially on aboveground carbon pools, studies 

that include eff ects of management activities on soil 

carbon have also been included. However, studies 

focused exclusively on soil carbon are not included in 

the scope of this bibliography. In some cases, studies 

with an explicit focus on net ecosystem productivity 

(NEP) or net primary productivity (NPP) have 

been included because the results include or draw 

conclusions about eff ects on carbon stocks.
3

Using this Bibliography

Th is annotated bibliography is available as a 

downloadable and printable document; the references 

included in the bibliography are also available as a 

downloadable EndnoteTM database or basic text fi le. 

For ease of use, the Appendix (beginning on page 44) 

provides summary tables containing all abbreviations 

of chemical elements and compounds, acronyms, tree 

scientifi c names used and corresponding common 

names, and units of measure and conversions used in 

this bibliography. Th is publication is available on our 

website at: http://www.nrs.fs.fed.us/pubs/. We are 

considering options for keeping this publication up-todate into the future; check back at the NRS website for 

the most recent updates.

Categories

I. Experiments and observations

Th is category contains results of lab and fi eld-based 

experiments and observation studies that explore 

the eff ects of a wide range of planned management 

activities and incidental or involuntary climate-related 

eff ects on carbon storage in natural and plantation 

forests.

II. Models

Th is wide-ranging category covers the diversity of 

models designed to forecast the potential eff ects of 

forest management decisions and/or changing climatic 

conditions and future scenarios and/or shifting and 

uncertain socioeconomic factors on carbon storage 

in forests, at the stand-level to the country-level. 

Th e category also includes studies that present 

carbon balance and fl ux data, forecasts, tradeoff s, or 

optimization results from models.

III. Experiments with some pools modeled

Th is category includes designed studies with fi eld 

plots. Th ese studies may report several years of 

experimental results and fi ndings, but rely on models 

to forecast future carbon storage. Studies that compare 

experimental and modeled results are also included in 

this category.

IV. Literature reviews and meta-analyses

Th is category includes broad-scale reviews and seminal 

synthesis papers which, by defi nition, rely on a variety 

of other sources to: 1) compile “big-picture” estimates 

of current or potential or both current and potential 

carbon storage by forests, 2) describe ways in which 

forest management, or land-use change, or both has 

or can aff ect carbon pools and cycling, or 3) describe 

likely eff ects of climate change and involuntary 

fertilization on forest health, productivity, and carbon 

cycling and storage.

V. Policy Perspectives

Th is category contains those studies that draw 

attention to considerations of importance to the 

creation, design, and implementation of policy 

to eff ectively mitigate climate change. Th e papers 

may be either explicit (contain specifi c policy 

recommendations) or may be more scientifi c in nature 

(highlighting uncertainties and considerations without 

suggesting policy measures), but because of their 

general or high-level nature are critical to informing 

forest carbon policy-related dialogue. In some cases, 

papers in this category may explicitly explore forest 

management decisions as described in “Experiments 

and observations” (category 1), but from a highlevel regional or national perspective. Th us, study 

conclusions are useful for broad-scale policy uses, 

rather than stand or forest-level management decisions.

LITERATURE CITED

Administrative Procedures Act of 1946 [APA]; 5 

U.S.C 1001-1011.

Forest and Rangeland Renewable Resources. Planning 

Act of 1974 [RPA]; 16 U.S.C 1601.

Heath, L.S; Kimble, J.M.; Birdsey, R.A.; Lal, R. 2003. 

Th e potential of U.S. forest soils to sequester 

carbon. In: Kimble, J. M.; Heath, L.S.; Birdsey, 

R.A.; Lal, R., eds.Th e potential of U.S. forest soils 

to sequester carbon and mitigate the greenhouse 

eff ect. Boca Raton, FL: CRC Publishing: 385-394.
4

Heath, L.S.; Smith, J.E. 2000. Soil carbon 

accounting and assumptions for forestry and 

forest-related land use changes. In: Joyce, L.A.; 

Birdsey, R., eds. Th e impact of climate change on 

America’s forests, Gen. Tech. Rep. RMRS-GTR-59. 

Fort Collins, CO: U.S. Department of Agriculture, 

Forest Service, Rocky Mountain Research Station: 

89-101.

Hoover, C.M. 2003. Soil carbon sequestration and 

forest management: challenges and opportunities.

In: Kimble, J.M.; Heath, L.S.; Birdsey, R.A.; 

Lal, R., eds. Th e potential of U.S. forest soils to 

sequester carbon and mitigate the greenhouse eff ect. 

Boca Raton, FL: CRC Publishing: 211-238.

Intergovernmental Panel on Climate Change. 2007. 

Climate change 2007: Synthesis report; Summary 

for policy makers. Available at http://www.jpcc.ch/

pdf/assessment-report/ar4/syr/ar4_syr_spm.pdf [23 

December 2008].

Johnson, D.W.; Curtis, P.S. 2001. Eff ects of forest 

management on soil C and N storage: meta 

analysis. Forest Ecology and Management.140: 

227-238.

Kimble, J.M.; Heath, L.S.; Birdsey, R.A.; Lal, R, 

eds. 2003. Th e potential of U.S. forest soils to 

sequester carbon and mitigate the greenhouse 

eff ect. New York: CRC Press. 448 p.

Nabuurs, G.J.; Masera, O.; Andrasko, K.; BenitezPonce, P.; Boer, R.; Dutschke, M.; Elsiddig, E.; 

Ford-Robertson, J.; Frumhoff , P.; Karjalainen, 

T.; Krankina, O.; Kurz, W.A.; Matsumoto, M.; 

Oyhantcabal, W.; Ravindranath, N.H.; Sanz 

Sanchez, M.J.; Zhang, X. 2007. Chapter 9: 

Forestry. In: Metz, B.; Davidson, O.R.; Bosch, 

P.R.; Dave, R.; Meyer, L.A., eds., Climate 

Change\2007: Mitigation. Contribution of working 

group III to the fourth assessment report of the 

Intergovernmental Panel on Climate Change. 

Cambridge, UK: Cambridge University Press. 

Available at: http://www.ipcc.ch/pdf/assessmentreport/ar4/wg3/ar4-wg3-chapter9.pdf [18 

December 2008].

National Forest Management Act of 2004 [NFMA]; 

Update of December 22, 2004. 36 CFR Part 219. 

U.S. Climate Change Science Program. 2008a. Th e 

fi rst state of the carbon cycle report (SOCCR): 

Th e North American carbon budget and 

implications for the global carbon cycle. In: King, 

A.W.; Dilling, L.; Zimmerman, G.P.; et al., eds. 

Asheville, NC: National Oceanic and Atmospheric 

Administration, National Climatic Data Center. 

242 p. Available at: http://www.climatescience.gov/

library/sap/sap2-2fi nal-report/sap2-2-fi nal-all.pdf 

[June 8, 2009].

U.S. Climate Change Science Program, 2008b. Th e 

eff ects of climate change on agriculture, land 

resources, water resources, and biodiversity in 

the United States. A report by the U.S. Climate 

Change Science Program and the subcommittee 

on global change research. Washington, DC: 

U.S Department of Agriculture. 240 p. Available 

at: http://www.usda.gov/oce/global_change/

fi lesCCSPFinalReport.pdf [June 8, 2009].

U.S. General Accounting Offi ce. 2003. Endangered 

species: Fish and Wildlife Service uses best 

available science to make listing decisions, but 

additional guidance needed for critical habitat 

designations. GAO-03-803. Washington, DC: 

U.S. General Accounting Offi ce. Available at: 

http://www.gao.gov/new.items/d03803.pdf [2 

February 2009].

U.S. Environmental Protection Agency. 2008.

Inventory of U.S. greenhouse gas emissions 

and sinks: 1990-2006. EPA 430-R-08-005. 

Washington, DC: U.S. Environmental Protection 

Agency, Offi ce of Atmospheric Programs. Available 

at: http://epa.gov/climatechange/emissions/U.S. 

gginv_archive.html [28 May 2009].
5

BIBLIOGRAPHY

I. Experiments and Observations: 

Management and Climate Effects

1. Adams, A.B.; Harrison, R.B.; Sletten, R.S.; 

Strahm, B.D.; Turnblom, E.C.; Jensen, C.M. 

2005. Nitrogen-fertilization impacts on carbon 

sequestration and fl ux in managed coastal 

Douglas-fi r stands of the Pacifi c Northwest.

Forest Ecology and Management. 220:313-325.

Th is study examines the eff ects of urea applications 

from1972 to 1996 on C sequestration in tree biomass 

and soils in Pseudotsuga menziesii for two soil types 

in western Washington. SOC was measured at four 

depths to a total of 1 m. Th ree to four applications 

of urea generally increased C density of both tree 

biomass and soils (average of 161 Mg C/ha vs. 135 

Mg C/ha for tree biomass on fertilized vs. unfertilized 

sites, respectively; average of 260 Mg C/ha vs. 175 

Mg C/ha for soil, respectively). Authors note that 

scope of study is limited due to small sample size and 

variation of fi eld installations. Because of this, they 

paired their data with Canary et al. (2001), analyzed 

for total C, and found an average increase of 82.8 

Mg C/ha for both biomass and soils at seven sites. 

Authors extrapolate over 10 million acres of intensively 

managed Pseudotsuga menziesii in the region, which 

yields an additional theoretical sequestration of 828 Tg 

C, but the study cautions that because all stands in the 

area are not of the same age and type, their assessment 

should not be widely applied.

2. Barford, C.C.; Wofsy, S.C.; Goulden, M.L.; 

Munger, J.W.; Pyle, E.H.; Urbanski, S.P.; Hutyra, 

L.; Saleska, S.R.; Fitzjarrald, D.; Moore, K. 

2001. Factors controlling long- and short-term 

sequestration of atmospheric CO2 in a midlatitude forest. Science. 294: 1688-1691.

Th is paper reports on 9 years of CO2 fl ux 

measurements for a 60- to 80-year-old northern 

hardwood forest (Harvard Forest) and compares results 

to biometric measurements of changes in live and dead 

wood pools for selected species. Findings from the fl ux 

study show net CO2 uptake of 2.0 +/- 0.4 Mg CO2/

ha/yr (0.55 +/- 0.11 Mg C/ha/yr) from 1993-2000, 

while the biometric measurements show a similar 

uptake of 1.6 +/- 0.4 Mg CO2/ha/yr (0.44 Mg C/ha/

yr +/- 0.11 Mg C/ha/yr).

Th e study reports that interannual carbon 

sequestration is aff ected in large part by previous 

management and the impacts of prior disturbances, 

whereas year-to-year fl uctuation in uptake is more a 

function of weather.

3. Balboa-Murias, M.Á.; Rodríguez-Soalleiro, 

R.; Merino, A.; Álvarez-González, J.G. 2006. 

Temporal variations and distribution of carbon 

stocks in aboveground biomass of radiata pine 

and maritime pine pure stands under diff erent 

silvicultural alternatives. Forest Ecology and 

Management. 237: 29-38.

Th is study examines the impact of fi ve site qualities, 

three stocking densities, and fi ve thinning intensities 

on aboveground biomass C sequestration in Pinus 

pinaster and Pinus radiata in plantation forests of 

northwest Spain, a region characterized by mild 

temperatures and a slight hydric defi ciency in summer. 

P. radiata has a higher NPP than P. pinaster, so 

total aboveground biomass C storage was greater for 

the former species. For P. radiata, mean annual C 

storage could be increased by 3.0 Mg/ha by choosing 

the highest stocking density, the best site quality, 

and the least intense thinning regime (15 percent). 

For the same scenario with maritime pine, annual 

C sequestration could be increased by 2.4 Mg/ha. 

Increasing the rotation age from 30-40 years yielded 

an increase in biomass of 33-38 percent, but mean 

annual C storage declined. Th is study concludes 

that sustainable management that seeks to optimize 

biomass production and C sequestration benefi ts most 

from good site quality, reduced thinning intensity, and 

extended rotation lengths.

4. Boerner, R.E.J.; Huang, J.; Hart, S.C. 2008. Fire, 

thinning, and the carbon economy: eff ects of 

fi re and fi re surrogate treatments on estimated 

carbon storage and sequestration rate. Forest 

Ecology and Management. 255: 3081-3097.

Th is study examines eff ects of prescribed fi re, 

mechanical treatments designed as surrogates 

for prescribed fi re, and combination treatments 
6

(mechanical plus fi re) on ecosystem C stocks at 12 

study sites in the eastern and western United States. 

Th ey found that while prescribed fi re had little impact 

on forest C budgets in eastern forests, mechanical 

or combination treatments converted the forests 

from net sinks to sources in the fi rst years following 

treatment, primarily due to a loss of aboveground 

biomass. Carbon budgets in western forests remained 

relatively stable, regardless of treatment. Overall, 

total ecosystem C across the network of sites was 

not signifi cantly aff ected by prescribed fi re, though 

several individual sites did experience signifi cant 

losses (p<.05); average net losses from mechanical 

or mechanical plus fi re ranged from 4-20 Mg/ha 

after 3 years (initial losses were often mitigated by 

increased uptake post-treatment), but only the losses 

in year 1 were signifi cant (p<.05). Findings suggest 

that optimal forest management for fi re, harvesting, 

and sequestration varies signifi cantly between eastern 

and western forests. Authors do not directly address 

to what extent treatments reduce risk of catastrophic 

wildfi re, and how this would aff ect C budgets, but do 

reference a variety of studies that indicate reduced U.S. 

forest sink-strength due to historical and continued fi re 

suppression.

5. Bond-Lamberty B.; Wang C.K.; Gower S.T. 2004. 

Net primary production and net ecosystem 

production of a boreal black spruce wildfi re 

chronosequence. Global Change Biology. 10: 473-

487.

Examines the C dynamics of a chronosequence of 

seven stands of Picea mariana of diff erent ages in the 

Canadian BOREAS Northern Study Area, west of 

Th ompson, Manitoba. Th e seven stands resulted from 

stand-replacing wildfi res around 1850 and in 1930, 

1964, 1981, 1989, 1995, and 1998. Th e NPP of 

stands in well and poorly drained sites were lowest in 

the years immediately following fi re (0.5-1.0 Mg C/

ha) and highest 12-20 years following fi re (3.32 in dry 

stands and 5.21 Mg C/ha in wet stands. NPP in the 

older stands was about half that in the 12-20 year old 

stands. For the entire chronosequence, NPP on dry 

sites ranged from 1.06-4.06 Mg C/ha, and NPP on 

wet sites ranged from 0.72-5.34 Mg C/ha. Th e study 

includes a summary table that includes NPP on dry 

and wet stands for each age according to aboveground 

and belowground measurements.

6. Bradford, J.B.; Birdsey, R.A.; Joyce, L.A.; Ryan, 

M.G. 2008. Tree age, disturbance history, and 

carbon stocks and fl uxes in subalpine Rocky 

Mountain forests. Global Change Biology. 14: 

2882-2897.

Th is study examines the relationship among tree age, 

history of stand disturbance, and forest C storage and 

cycling on seven stands at three sites to determine 

whether either age, disturbance history, or both 

can be used to predict C stocks and fl uxes for three 

distinct subalpine central Rocky Mountain forest 

ecosystems (Fraser Experimental Forest, Glacier Lakes 

Ecosystem Experimental Site (GLEES), Niwot Ridge 

Amerifl ux Study Site). Th e sites diff ered in tree age, 

disturbance history, species composition, elevation, 

and precipitation, but ecosystem C storage was similar 

(roughly 260 Mg C/ha at Fraser and Niwot and 330 

Mg C/ha at GLEES), with 80-90 percent of live C 

stored in trees. “Maximum tree age overestimated time 

since disturbance for young forests and underestimated 

it for old forests,” making tree age a poor predictor of 

disturbance history. Tree age and disturbance history 

infl uenced both C pools and fl uxes, but there is little 

change after the fi rst 100 years has passed following a 

disturbance. Because of this, the study concludes that 

landscape-scale estimates can be simplifi ed by treating 

subalpine forests in this region as a single forest type 

and that the 100 years following a disturbance is the 

most critical factor aff ecting C change.

7. Canary, J.D.; Harrison, R.B.; Compton, J.E.; 

Chappell, H.N. 2000. Additional carbon 

sequestration following repeated urea 

fertilization of second-growth Douglas-fi r stands 

in western Washington. Forest Ecology and 

Management. 138: 225-232.

Eff ects of urea application over a 16-year period are 

examined for Pseudotsuga menziesii stands (age 62-

69 at study’s end) in western Washington. Average 

aboveground tree C increased by 26.7 Mg C/ha over 

the control during the 16-year period (signifi cant 

at p<.01), or 1.7 Mg C/ha on an annualized basis 
7

(though increases likely were not constant). Total 

ecosystem C increased by 34.8 Mg C/ha over the 

control (2.2 Mg/ha on an annualized basis, though 

increases were likely not constant), but diff erences in 

individual snag, soil, and understory pools were not 

signifi cant. Th e fi ndings emphasize the importance of 

tracking soil C below the surface horizons; 75 percent 

of soil C (to a depth of 85 cm) was found below the 

A-horizon.

8. Caspersen J.P.; Pacala, S.W.; Jenkins, J.C.; Hurtt, 

G.C.; Moorcroft, P.R.; Birdsey, R.A. 2000. 

Contributions of land-use history to carbon 

accumulation in U.S. forests. Science. 290: 1148-

1151.

Th e study examines the relative importance of landuse history and CO2 fertilization, N deposition, 

and climate change on the C uptake of forests in 

Minnesota, Michigan, Virginia, North Carolina, and 

Florida. Comparison FIA data (includes estimates of 

aboveground biomass, growth- and mortality-related 

changes in aboveground biomass, and stand age) 

from the late 1970s/early 80s to the mid 1990s were 

used for 20,000 plots; results indicate that growth 

enhancement (defi ned as CO2 fertilization and N 

deposition) accounted for only 2.0 +/- 4.4 percent of 

aboveground NEP.

9. Chiang, J.-M.; McEwan, R.W.; Yaussy, D.A.; 

Brown, K.J. 2008. Th e eff ects of prescribed fi re 

and silvicultural thinning on the aboveground 

carbon stocks and net primary production of 

overstory trees in an oak-hickory ecosystem in 

southern Ohio. Forest Ecology and Management. 

255: 1584-1594.

Th is study examines the impacts of thinning and 

prescribed burns on Quercus recruitment and 

regeneration, and assesses changes in C storage and 

ANPP due to thinning/burning treatments. Estimated 

pretreatment aboveground biomass was 233 Mg/

ha, and thinning resulted in a 30 percent reduction 

in aboveground biomass. Authors did not trace the 

carbon lost from thinning practices to determine how 

much was ultimately retained in long-term wood 

products, but did fi nd a substantial but transient 

reduction in ANPP in thinned stands for both fi eld 

measurements and ancillary model simulations that 

directly replicated the fi eld experiment. Prescribed 

burning has a signifi cant impact (p<.05) on the 

mortality of stems (of both Acer and Quercus; the latter 

was less fi re-resistant than is typically expected) but did 

not signifi cantly aff ect ANPP. Neither thinning nor 

burning has a long-term impact on C uptake in these 

forests.

10. Coyle, D.R.; Coleman, M.D. 2005. Forest 

production responses to irrigation and 

fertilization are not explained by shifts in

allocation. Forest Ecology and Management. 208: 

137-152.

Th is study examines aboveground and belowground 

growth and biomass accumulation response of two 

Populus deltoides clones and Platanus occidentalis

that received irrigation (I), fertilization (F), or a 

combination treatment (IF) in South Carolina. Total 

biomass (branch, bark, wood, stump, coarse and fi ne 

root) response of nearly every tree to both I and F 

treatments (40, 80, 120 kg N/ha/yr for years 1, 2, 

3, respectively) was signifi cant (p< .05). Trees that 

received the IF treatment accumulated signifi cantly 

more biomass than those in other treatments (p<.05). 

P. occidentalis that received the IF treatment had the 

most signifi cant gain in biomass C gain, with a mean 

annual increase of 6.8 Mg C/ha/yr; combination 

treatments for the P. deltoides clones yielded increases 

of 3.9 and 3.5 Mg C/ha/yr. Th ese was no signifi cant 

diff erence between root and shoot allocation.

11. Curtis, P.S.; Hanson, P.J.; Bolstad, P.; Barford, C.; 

Randolf, J.C.; Schmid, H.P.; Wilson, K.B. 2002. 

Biometric and eddy-covariance based estimates 

of annual carbon storage in fi ve North American 

deciduous forests. Agricultural and Forest 

Meteorology. 113: 3-19.

Th is study examines and compares annual C storage 

for fi ve deciduous U.S. forest sites of similar age, 

canopy height, and stand basal area across Tennessee, 

Indiana, Massachusetts, Michigan, and Wisconsin. 

Meteorological and biometric measurements of 

C storage were compared over time. Biometric 
8

measurements indicated an NEP of between 0.7-3.5 

Mg C/ha/yr among the sites (highest in south-central 

Indiana, lowest in upper Michigan), with an average 

of 1.9 Mg C/ha/yr, while the meteorological model 

results yielded values of about 3.0-5.0 Mg C/ha/yr 

for aboveground net primary productivity (the higher 

values corresponded to more southern sites). Th e study 

states that diff erences in biometric NPP estimates 

correlated to expectations based on site-specifi c 

soil and climate diff erences. Th ere were no major 

diff erences between biometric and meteorological 

estimates for same-site comparisons, and diff erences 

were smaller when “fewer belowground processes were 

explicitly incorporated into the estimates.”

12. Finkral, A.J.; Evans, A.M. 2008. Th e eff ects 

of a thinning treatment on carbon stocks in a 

northern Arizona ponderosa pine forest. Forest 

Ecology and Management. 255: 2743-2750.

Th e study examines the C consequences of restoration 

thinning practices designed to reduce wildfi re risk on 

the aboveground C balance of a Pinus ponderosa forest 

in Arizona. Treatment reduces risk of catastrophic fi re, 

but it still results in a one-time net loss of 3.11 Mg C/

ha. Simply using the slash to create long-lived wood 

products (rather than burning) results in C benefi ts 

of 3.35 Mg C/ha. Th is fi nding that has important 

implications for hazardous fuels treatments throughout 

the western United States.

13. Fitzsimmons, M.J.; Pennock, D.J.; Th orpe, J. 

2004. Eff ects of deforestation on ecosystem 

carbon densities in central Saskatchewan, 

Canada. Forest Ecology and Management. 188: 

349-361.

Findings from Saskatchewan, Canada indicate that 

forests store nearly twice as much C as compared to 

cultivated fi elds (158 vs. 81 Mg C/ha, respectively). 

Conversion of forest to agricultural land would 

result in near-term C losses of at least 30 Mg C/ha. 

Reforestation of pasture and agricultural land with 

trembling aspen could sequester 30-75 Mg C/ha over a 

50- to 100-year time horizon.

14. Foster, N.W.; Morrison, I.K. 2002. Carbon 

sequestration by a jack pine stand following urea 

application. Forest Ecology and Management. 169: 

45-52.

Th e study examines C accumulation response of semimature Pinus banksiana stands in Ontario, Canada 

to the application of a one-time N amendment of 0 

(control), 56, 112, 224, or 448 kg N/ha. Th ere is no 

signifi cant diff erence in the long-term (30-year) net 

C accumulation of the stands. However, there is a 

statistically signifi cant increase (p<.05) in diameter at 

breast height (d.b.h.) increment over the codominant 

species at year 10, suggesting that urea application 

10 years before harvest can increase timber yield. Th e 

study concludes that over-stocked, semi-mature P. 

banksiana stands should be given low priority when 

considering nutrient amendments, but states that C 

accumulation may have been constrained by other 

nutrients and a combination N/P/K amendment 

may lead to substantial long-term aboveground 

accumulation of C.

15. Giasson, M.-A.; Coursolle, C.; Margolis, HA. 

2006. Ecosystem-level CO2 fl uxes from a boreal 

cutover in eastern Canada before and after 

scarifi cation. Agricultural and Forest Meteorology. 

140: 23-40.

Th is study uses CO2 fl ux towers and the eddy 

covariance technique to study the eff ects of 

scarifi cation on NEP of a recently harvested boreal

Picea mariana/Pinus banksiana stand in Quebec, 

Canada. Scarifi cation was necessary to produce 

adequate regeneration, yet the treatment, which 

destroyed 60 percent of the aboveground biomass, 

resulted in the site becoming a notably greater C 

source following treatment (1.11 Mg C/ha before vs. 

1.75 Mg C/ha after treatment). Scarifi cation resulted 

in biomass destruction and daily NEP was lower for 

the year following treatment. Th e fi ndings suggest 

that cutovers may account for emissions of 5 Mt C/

yr in Canada (generalized to all Canadian boreal 

ecosystems. Th is study does not make mention of 

whether or to what extent scarifi cation eventually (year 

5 and beyond) leads to greater C uptake in regenerated 

stands.
9

16. Gough, C.M.; Vogel, C.S.; Harrold, K.H.; 

George, K.; Curtis, P.S. 2007. Th e legacy of 

harvest and fi re on ecosystem carbon storage in 

a north temperate forest. Global Change Biology. 

13: 1935-1949.

Th is study examines the eff ects of disturbances 

(clearcut and subsequent burn) on C storage in 

a mixed deciduous forest in the northern Lower 

Peninsula of Michigan. Although stands that were 

repeatedly disturbed reverted to sinks within 6 

years of disturbance, the long-term trajectory and 

overall capacity of C storage was limited. Repeated 

disturbances (a second harvest and fi re) reduced site 

quality and resulted in annual C storage that was 

45 percent lower than that in the reference stand 

receiving a single harvest-fi re disturbance. Maximum 

annual C storage that was 26 percent lower than that 

in the reference stand over a 62-year period. Previous 

harvesting regimens that reduced site quality continue 

to limit C sequestration in these areas.

17. Gough, C.M.; Vogel, C.S.; Schmid, H.P.; Curtis, 

P.S. 2008. Controls on annual forest carbon 

storage: lessons from the past and predictions for 

the future. BioScience. 58: 609-622.

In this study, authors use a combination of ecological 

and meteorological approaches to measure annual 

forest C storage from 1999-2005 in a mixed deciduous 

forest in Michigan’s northern Lower Penninsula. 

Th e area is representative of 102,000 km2 of forest 

in the upper Midwest that is dominated by Populus 

grandidentata and Populus tremuloides. Th e study 

examines the response of annual C storage to historical 

disturbances, and fi nds that repeated clearcut harvests 

followed by fi re result in lower overall ecosystem C 

storage capacity. In this ecosystem (average stand age 

85 years, with stands ranging in age from 6-90 years), 

net C gain was about 1.5 Mg C/ha/yr, a rate typical of 

other aspen-dominated forests. Th e study concludes: 

“Eff ective forest C sequestration requires management 

of all C pools, including traditionally managed pools 

such as bole wood and also harvest residues and soils.”

18. Guo, L.B.; Bek, E.; Giff ord, R.M. 2006. Woody 

debris in a 16-year-old Pinus radiata plantation 

in Australia: Mass, carbon and nitrogen stocks, 

and turnover. Forest Ecology and Management. 

228: 145-151.

Authors examine the C density of woody debris on 

aff orested (with Pinus radiata) former pastureland 

in Australia. A previous meta-analysis spanning 

temperate, tropical, and subtropical forests in 16 

countries demonstrated that soil C stocks decreased 

following conversion of native pasture to forest 

plantation (Guo and Giff ord 2002). Th is study 

hypothesizes that soil C loss from land-use change is at 

least partially off set by gains in the woody debris pool. 

Woody debris contained 3.1 Mg C/ha in a 16-yearold Pinus radiata plantation 8 years after thinning and 

pruning activities, which represents approximately 

22 percent of soil carbon reduction from previous 

land-use change. Th e pine plantation was a net C sink 

because gains in aboveground biomass exceeded SOC 

losses. Th e decay of woody debris is infl uenced by 

resource quality and environmental conditions such 

as temperature, moisture, aeration, and decomposer 

organisms. Th e publication contains extensive data on 

decomposition and C release rate for various species 

and diameter classes of pine.

19. Harmon, M.E.; Bible, K.; Ryan, M.G.; Shaw, 

D.C.; Chen, H.; Klopatek, J.; Li, X. 2004. 

Production, respiration, and overall carbon 

balance in an old-growth Pseudotsuga-Tsuga

forest ecosystem. Ecosystems. 7: 498-512.

Th is study uses ground-based measurements to 

determine the GPP, NPP, NEP, and autotrophic and 

heterotrophic respiration of an old-growth PseudotsugaTsuga forest at the Wind River Experimental Forest 

in Washington. Th e study reports average values for 

live C stores (39.8 Mg C/ha), detritus and mineral 

soil (22.1 Mg C/ha) and total C (61.9 Mg C/ha), and 

fi nds a weak sink of approximately 0.2 Mg C/ha/yr. 

Th ese results diff er from estimates using an eddy-fl ux 

method but the diff erence could be the result of a 

temporal mismatch between datasets.

20. Hicke, J.A.; Asner, G.P.; Kasischke, E.S.; French, 

N.H.F.; Randerson, J.T.; Collatz, G.J.; Stocks, 

B.J.; Tucker, C.J.; Los, S.O.; Field, C.B. 2003. 
10

Postfi re response of North American boreal forest 

net primary productivity analyzed with satellite 

observations. Global Change Biology. 9: 1145-

1157.

Th is study uses a 17-year record of satellite NDVI 

observations to identify fi re scars across the North 

American boreal landscape and determine NPP in 

post-burn years in these fi re-scarred areas using the 

CASA carbon-cycle model. NPP decreased by an 

average of 0.6-2.6 Mg C/ha (30-80 percent). On 

average, recovery period for boreal forests is about 9 

years, though results vary substantially among fi res.

21. Howard, E.A.; Gower, S.T.; Foley, J.A.; Kucharik, 

C.J. 2004. Eff ects of logging on carbon dynamics 

of a jack pine forest in Saskatchewan. Canada. 

Global Change Biology. 10: 1267-1284.

Th is study measures the NPP of a chronosequence of 

fi ve Pinus banksiana stands in Saskatchewan, Canada 

that was part of the southern area of the BOREAS 

study site. Th e four younger stands (0, 5, 10, and 29 

years old) had been harvested; the oldest (79 years 

old) originated following wildfi re. Average NPP 

measurements on each stand were positive: 0.9, 1.3, 

2.7, 3.5, and 1.7 Mg C/ha, respectively, but NEP, 

which takes into account heterotrotrophic respiration 

by soils, was negative for the two youngest and the 

oldest stand, and only slightly positive (0.4) for the 10- 

and 29-year-old stands. Spatial and methodological 

uncertainties limit conclusions, but results show that 

stands become an atmospheric C source in the years 

following logging.

22. Hubbard, R.M.; Vose, J.M.; Clinton, B.D.; 

Elliott, K.J.; Knoepp, J.D. 2004. Stand restoration 

burning in oak-pine forests in the southern 

Appalachians: eff ects on aboveground biomass 

and carbon and nitrogen cycling. Forest Ecology 

and Management. 190: 311-321.

Th e study considers the eff ects of understory prescribed 

burning on forest biomass and C pools on oakdominated forest stands with fi ne-loamy, mixed, mesic 

soils in southern Tennessee and Georgia. Th ere was 

no signifi cant diff erence (p>0.1) in total aboveground 

live woody biomass between burned and control plots. 

However, in burned plots, understory biomass alone 

was reduced by 50 percent (p=.07) and overstory 

biomass was reduced by 11 percent, but the diff erence 

in the latter case was not signifi cant. Burning 

signifi cantly reduced (p<.01) C pools in CWD, small 

wood, and litter. Th e total forest fl oor (CWD, small 

wood, litter, and humus) C pool was reduced by 4.05 

Mg C/ha, but this diff erence was also not signifi cant 

(p> 0.1). C fl ux was reduced on burned sites for 2 

months following treatment, but then returned to 

normal. Monthly litterfall was not aff ected, suggesting 

that pools would be restored quickly. Prescribed 

burning can be an eff ective management tool that does 

not adversely aff ect C cycling or nutrient cycling.

23. Hazlett, P.W.; Gordon, A.M.; Sibley, P.K.; Buttle, 

J.M. 2005. Stand carbon stocks and soil carbon 

and nitrogen storage for riparian and upland 

forests of boreal lakes in northeastern Ontario. 

Forest Ecology and Management. 219: 56-68.

Research in northeastern Ontario, Canada to 

determine how forest management and harvesting 

activities may impact ecosystem C levels examined 

C and N storage in upland and riparian boreal forest 

ecosystems. Total, aboveground, and belowground 

C stocks did not diff er signifi cantly (p>.05) between 

upland and riparian stands (total C was 83.847 Mg C/

ha riparian vs. 81.487 Mg C/ha upslope). However, in 

the riparian zones, a greater proportion of total C was 

stored in overstory stocks and fl oor layers were deeper 

and contained more C than upland sites. Because of 

these diff erences in C allocation between zones, similar 

harvesting techniques (whole-tree harvesting) removes 

far more aboveground and total C in riparian than 

upland boreal ecosystems (98 percent vs. 76 percent, 

respectively for aboveground; 35 percent vs. 17 percent 

respectively, for total ecosystem C). Because riparian 

ecosystems and the aquatic habitats they support are 

particularly sensitive to changes in nutrient levels, 

changes in harvesting techniques (selective, partial, 

modifi ed) would result in greater C retention in these 

ecosystems, and likely faster postharvest regeneration 

due to greater nutrient availability.

24. Laclau, P. 2003. Biomass and carbon 

sequestration of ponderosa pine plantationsand 
11

native cypress forests in northwest Patagonia. 

Forest Ecology and Management. 180: 317-333.

Conversion of pasture to pine plantations in northwest 

Patagonia resulted in little change in soil C but 

created an additional biomass sink of 3.5 Mg C/ha 

during the plantations’ fi rst 15 years. In signifi cantly 

older but uneven-aged cypress stands that had been 

subject to intense grazing during previous harvests 

there was no signifi cant diff erence in total ecosystem 

C from pine plantations, but signifi cantly greater soil 

C (an additional 20 Mg C/ha more in cypress stands, 

to a depth of 50 cm). ANOVA revealed signifi cant 

diff erences in soil C between forest types but not 

between land uses (forest vs. pasture). Converting 

cypress forests to pine plantations could result in 

substantial losses in soil C. Because cypress and pine 

grow on distinctly diff erent soil types in the United 

States, and conversion from one forest type to the 

other may not be feasible, and conversion of pasture to 

forest should exceed the baseline pasture sequestration 

rate of 2.6 Mg C/ha.

25. Laiho, R.; Sanchez, F.; Tiarks, A.; Dougherty, 

P.M.; Trettin, C.C. 2003. Impacts of intensive 

forestry on early rotation trends in site carbon 

pools in the southeastern U.S. Forest Ecology and 

Management. 174: 177-189.

Intensive management of southern forest plantations 

has led to threefold increases in productivity, but 

management and harvesting eff ects on C pools is 

not completely understood. Th is study examines 

the eff ects of four harvesting strategies (stem only, 

whole tree, whole tree with forest fl oor removal, and 

full amelioration, consisting of whole tree, disking, 

bedding, and fertilization), in some cases combined 

with herbicide treatments, on C pools in managed 

forest plantations in North Carolina and Louisiana. 

Th e plantations consisted of 55-year-old mixed pinehardwood stands, with Pinus taeda as the dominant 

species. Measurements were taken prior to harvest and 

5 years following harvest. Th ere were no substantial 

changes in the upper soil C profi le (<30 cm) in the 

short term by treatment, but there were only three 

soil samples per plot on the North Carolina site. 

Site preparation may have positively infl uenced soil 

C pools. On the North Carolina site, the amount 

of organic matter removed played a larger role than 

herbicide treatment in infl uencing soil C; the opposite 

was true for the Louisiana site. Aboveground and 

soil C pools depended on both harvesting and site 

preparation methods and levels of signifi cance varied 

among sites. Whole-tree harvesting reduced soil C 

(p<.03), amelioration and fertilization increased soil C 

in North Carolina (p<.06), herbicides reduced soil C 

in all cases in Louisiana, all treatments reduced forest 

fl oor C in North Carolina (p<.06), and whole-tree 

harvesting with or without herbicides reduced forestfl oor C in Louisiana (p<.06).

26. Lee, J.; Morrison, I.K.; Leblanc, J.-D.; Dumas, 

M.T.; Cameron, D.A. 2002. Carbon sequestration 

in trees and regrowth vegetation as aff ected by 

clearcut and partial cut harvesting in a secondgrowth boreal mixed wood. Forest Ecology and 

Management. 169: 83-101.

Th e study examines the impacts of forest management 

practices (three silvicultural options: clearcut vs. 

shelterwood vs. uncut control; and two harvesting 

systems: tree-length vs. whole-tree removal) and 

post-harvest ecosystem C pools and recovery in 

boreal mixed-wood forests in Ontario, Canada. 

Dominant species were Picea glauca, Picea mariana, 

Abies balsamea, Populus tremuloides, and Betula alba. 

Post-harvest C assimilation rates for the clearcut and 

partial cut plots were similar and signifi cantly greater 

(p<.001) than those for the control (uncut) plot. Also, 

total ecosystem sequestration was signifi cantly greater 

(p<.001) on control than on partial vs. clearcut plots 

(roughly 3.1 Mg C/ha/yr vs. 1.8 Mg C/ha/yr vs. 0.3 

Mg C/ha/yr) 5 years after harvest. Results demonstrate 

the rapid regrowth of ground vegetation followed by 

aspen thickets, which speeded the reconstitution of 

ecosystem C pools. C sequestration on partial-cut plots 

would have been closer to that of control plots had 

windthrow not occurred and more care been taken 

to minimize damage during harvesting operations. 

Although partial harvesting results in greater ecosystem 

C assimilation than clearcutting, this must be weighed 

against “lower productivity and higher costs” that 

result from partial cutting. Th is study did not account 

for carbon in harvested wood products.
12

27. Li, Q.; Chen, J.; Moorhead, D.L.; DeForest, 

J.L.; Jensen, R.; Henderson, R. 2007. Eff ects of 

timber harvest on carbon pools in Ozark forests.

Canadian Journal of Forest Research. 37: 2337-

2348.

Th is study compares the eff ects of no-harvest 

management, single-tree uneven-age management, 

and clearcut even-age management on forest C 

pools 8 years after treatment in the Missouri Ozarks. 

Dominant species were Quercus alba, Quercus velutina, 

Quercus coccinea, Carya glabra, and Carya cordiformis. 

Although ecosystem C (live tree, roots, CWD, 

forest-fl oor litter, and soil to a depth of 15 cm) was 

greatest for the no-harvest treatment (182 Mg C/

ha), the single-tree uneven-age treatment contained 

172 Mg C/ha and resulted in a 31 percent loss of C 

in the live-tree pool, compared with 130 Mg C/ha 

and a 93 percent decline in the live-tree C pool in the 

clearcut even-aged management treatment. However, 

diff erences in ecosystem C were signifi cant (p<.05) 

for each treatment. Eff orts to increase ecosystem C 

sequestration must consider C pool allocation patterns 

of specifi c ecosystems, and how management might 

reconfi gure these pools. Th is study did not account for 

carbon in harvested wood products.

28. Litton, C.M.; Ryan, M.G.; Knight, D.H. 2004. 

Eff ects of tree density and stand age on carbon 

allocation patterns in postfi re lodgepole pine. 

Ecological Applications. 14: 460-475.

Th e study examines how post fi re C pools, fl uxes, and 

allocation patterns varied among 13- and 110-yearold stands of Pinus contorta in Yellowstone National 

Park. Th e stands diff ered in tree density; C in biomass 

and detritus was estimated from direct sampling and 

allometric equations. Most of the C (91-99 percent) 

in young stands was detritus; in mature stands, 64 

percent of the C was in live vegetation. In young 

stands, C (foliage, stems, coarse and fi ne roots) 

increased signifi cantly (p<.01) with tree density and 

C in live vegetation increased signifi cantly (p<.01) 

as stands matured. Total ecosystem C in each stand 

(13-year-old low, moderate, and high density stands, 

and 110-year-old mature stand) was 76.36, 70.00, 

63.70, and 170.79 Mg C/ha, respectively. Th ere 

was a constant ratio of ANPP and TBCA over time, 

implying that C allocation patterns in P. contorta are 

independent of stand age or tree density.

29. McCarthy, H.R.; Oren, R.; Kim, H.S.; Johnsen, 

K.H.; Maier, C.; Pritchard, S.G.; Davis, M.A.

2006. Interaction of ice storms and management 

practices on current carbon sequestration in 

forests with potential mitigation under future 

CO2 atmosphere. Journal of Geophysical Research. 

111: D15103.

Few estimates of future sequestration and C budgets 

account for the eff ects of nonfi re disturbance on 

C sequestration and release. Th is study emphasizes 

the importance of considering the eff ects of ice 

storms, which “aff ect forest ecosystems on a range of 

timescales” (immediate tree mortality, reduced future 

productivity for damaged trees, reduced reproductive 

capacity for some species that produce seeds in the 

upper crown) and occur approximately once every 

6 years in the southeastern United States. Practices 

designed to increase wood quality and/or yield, such 

as thinning and/or fertilization, can initially leave 

forests more vulnerable to damage from ice storms. (A 

recently thinned stand has low stand density and no 

crown-to-crown support, and it has been shown that 

fertilization reduced wood density and strength.) Th e 

eff ects of ice-storm damage were examined on thinned, 

fertilized, and control plots in both current and 

elevated CO2 conditions (the latter through a FACE 

site). Fertilization resulted in no signifi cant diff erence 

(p>.10), but thinned plots under current climate 

conditions suff ered signifi cantly greater damage than 

unthinned plots. Both types of plots incurred about 60 

percent less damage under elevated CO2 conditions. 

Th e study concludes that loblolly pine plantations, 

currently restricted by temperature and ice-storm 

constraints, could be expanded northward because they 

would experience less damage and mortality under 

elevated CO2 conditions.

30. Morehouse, K.; Johns, T.; Kaye, J.; Kaye, M. 

2008. Carbon and nitrogen cycling immediately 

following bark beetle outbreaks in southwestern 

ponderosa pine forests. Forest Ecology and 

Management. 255: 2698-2708.
13

Th is study examines the eff ects of bark beetle 

infestation on C and N pools and fl uxes in stands of 

Pinus ponderosa in Arizona. Bark beetles have been 

a perennial source of low-level disturbance in pine 

forests, but this paper states that their impact has been 

magnifi ed by drought and fi re exclusion (it does not 

cite climate change as an exacerbating factor.) On sites 

with similar ecosystem structure before infestation, 

there were signifi cant diff erences (p<.05) between 

infested and uninfested sites in live- and dead-tree 

C pools. Infested sites held 65 Mg C/ha (60 percent 

of aboveground biomass) in dead trees compared to 

12 Mg C/ha (5 percent) in uninfested plots. Th ere 

were signifi cant diff erences in needle N-concentration 

(which was higher in infested plots due to premature 

needlefall) 3 years after infestation, but no diff erences 

in soil respiration rates between sites. Th is study notes 

that additional research is needed to better understand 

long-term C fl ux and storage impacts that result from 

signifi cant tree mortality.

31. Mund, M.; Kummetz, E.; Hein, M.; Bauer, G.A.; 

Schulze, E.-D. 2002. Growth and carbon stocks 

of a spruce forest chronosequence in central 

Europe. Forest Ecology and Management. 171: 

275-296.

Th e study attempts to isolate indirect and direct 

human impacts on forest growth in six monotypic 

Picea abies stands in Bavaria, Germany. Each stand 

was a diff erent age (16, 35, 43, 72, 112, and 142 

years old) but ecosystem conditions (acidic soils, 

high N deposition) were similar. Actual growth 

(independent of age) at both the tree and stand level 

exceeded that predicted by growth and yield tables, 

but growth was stimulated due to thinning and forest 

management. Carbon sink capacity in the future 

cannot be predicted by historical growth and yield 

tables based on undisturbed site conditions. Th e causes 

of altered growth remain unclear but might be related 

to increased CO2 concentrations, N deposition, 

elevated temperature, and the eff ects of thinning and 

management (or combined synergistic eff ects).

32. Nilsen, P.; Strand, L.T. 2008. Th inning 

intensity eff ects on carbon and nitrogen stores 

and fl uxes in a Norway spruce (Picea abies (L.) 

Karst.) stand after 33 years. Forest Ecology and 

Management. 256: 201-208.

Th is study presents results of a long-term fi eld 

experiment to evaluate the eff ects of thinning 

treatments on a then 22-year-old Picea abies stand 

in Norway. Th e stand was thinned from 3,190 to 

2,070, 1,100, and 820 trees/ha, respectively; this 

treatment was replicated three times to create four 

plots of each treatment. Results from 32-33 years 

after thinning indicate a signifi cant decrease (p<.05) 

in standing biomass C storage (trunk, bark, living 

and dead branches, stumps, and coarse and fi ne roots) 

for the N1100 stand and the N870 stand (22 and 

27 percent decrease, respectively) compared to the 

N2070 treatment (diff erence between the N1100 

and N870 stands was not signifi cant). Th inning 

generally is thought to increase volume production 

of remaining trees in the stand, but this was not true 

in this experiment, the results of which correspond 

to similar fi ndings in other experiments in Norway. 

Diff erences in ground vegetation between thinning 

treatments were not signifi cant (p>.05), but there was 

large variation between plots. Th ere was a negative 

correlation between standing volume and C storage 

in ground vegetation. Litterfall C was lower on more 

intensively thinned plots but the diff erence was not 

signifi cant. Th e study concludes that 32-33 years is 

likely too short a period in which to detect diff erences 

in soil C pools based on thinning treatments. 

However, longer time periods may not reveal greater 

diff erences given the variability and sampling scheme.

33. Shan, J.; Morris, L.A.; Hendrick, R.L. 2001. Th e

eff ects of management on soil and plant carbon 

sequestration in slash pine plantations. Journal of 

Applied Ecology. 38: 932-941.

Th is study examines the eff ects of three management 

treatments (fertilization, understory elimination, and 

fertilization plus understory elimination) and a control 

on aboveground and soil biomass of a 17-year-old 

Pinus elliottii stand in northern Florida. All treatments 

signifi cantly (p<.05) increased tree biomass and thus 

tree C (stems, branches, and needles). Eliminating 

the understory increases forest fl oor biomass, but 

results in lower soil C storage for all three treatments 
14

(p<.05), though not all diff erences among treatments 

were signifi cant. Extensive C pool tables are included 

in this publication. Th is study concludes that 

intensive management can increase sequestration in 

aboveground biomass and subsequent wood products, 

though soil C storage remains unaff ected or may 

decrease in some cases.

34. Smithwick, E.A.H.; Harmon, M.E.; Remillard, 

S.M.; Acker, S.A.; Franklin, J.F. 2002. Potential 

upper bounds of carbon stores in forests of the 

Pacifi c Northwest. Ecological Applications. 12: 

1303-1317.

Old growth forests are signifi cant C stores particularly 

in the highly productive Pacifi c Northwest. Th is study 

examines 43 old growth stands (150-1200 years old, 

without a major disturbance) in fi ve distinct biomes to 

determine the upper bound of C sequestration in the 

region, absent anthropogenic and natural disturbances. 

Coastal Oregon stands represent the upper bound and 

can accumulate stocks of up to 1,127 Mg C/ha (total 

ecosystem C); average area-weighted total ecosystem C 

for the Pacifi c Northwest was 671 Mg C/ha to a soil 

depth of 1.0 m, and 640 Mg C/ha to a soil depth of 50 

cm.

35. Sullivan, B.W.; Kolb, T.E.; Hart, S.C.; Kaye, 

J.P.; Dore, S.; Montes-Helu, M. 2008. Th inning 

reduces soil carbon dioxide but not methane fl ux 

from southwestern U.S.A. ponderosa pine forests. 

Forest Ecology and Management. 255: 4047-4055.

Th is study examines the eff ects of a thinning treatment 

on CO2 and methane fl uxes, forest-fl oor and understory 

biomass, and soil temperature, water content, and 

microbial biomass C in a Pinus ponderosa stand in 

Arizona. Th inning reduces soil CO2 effl ux immediately 

after treatment but has no eff ect on methane oxidation 

rates during the post-treatment months measured. 

Forest-fl oor mass at each site was statistically similar 

(p=.899) 2 years after thinning; aboveground understory 

plant biomass was signifi cantly greater (p<.05) at the 

thinned site after 1 year.

36. Van Miegroet, H.; Moore, P.T.; Tewksbury, 

C.E.; Nicholas, N.S. 2007. Carbon sources 

and sinks in high-elevation spruce-fi r forests 

of the Southeastern U.S. Forest Ecology and 

Management. 238: 249-260.

Th e study measures C pools and fl uxes in a mature 

high-elevation Picea rubens-Abies fraser forest in the 

Great Smoky Mountains National Park, on the border 

of North Carolina/ Tennessee. Although aboveground 

spruce C pools are highly variable, average C storage 

is estimated at 403 Mg C/ha. More than half of this 

storage was in soils whereas dead wood (mostly fi r) 

accounted for as much as 15 percent of ecosystem C. 

Th e study estimates mean live tree sequestration (2.7 

Mg C/ha/yr), dead wood release (1.6 Mg C/ha/yr) 

and net soil effl ux (1-1.45 Mg C/ha/yr), depending 

on assumptions, and concludes that the system is near 

carbon-neutral.

37. Vesterdal, L.; Rosenqvist, L.; Salm, C.; Hansen,K; 

Groenenberg, B.J.; Johansson, M.B. 2007. Chapter 

2: Carbon sequestration in soil and biomass 

following aff orestation: experiences from oak and 

Norway spruce chronosequences in Denmark, 

Sweden and the Netherlands. In: Heil, G.W.; 

Muys, B.; Hansen, K., eds. Environmental eff ects of 

aff orestation in North- Western Europe. Dordrecht, 

Th e Netherlands: Springer: 19-51.

Th is study examines C pools associated with six 

common European tree species 30 years after 

aff orestation. Th ere were pronounced diff erences in 

the forest-fl oor C pool and, in some cases, diff erences 

in the soil C pool. Th ere were no major diff erence 

in the combined forest-fl oor and mineral-soil layers, 

though species with smaller forest-fl oor C pools tended 

to have larger soil C pools. Th ere were diff erences in 

storage profi les (but not necessarily total storage) of 

C, N, and C/N ratios within the fi ve deciduous tree 

species. Forest-fl oor pools were greatest for Picea abies, 

followed by Fagus spp. and Quercus spp.; Fraxinus, 

Tilia, and Acer had the least C accumulation in forest 

fl oor pools.

38. Wang, C.; Bond-Lamberty, B.; Gower, S.T. 

2003. Carbon distribution of a well- and poorlydrained black spruce fi re chronosequence. Global 

Change Biology. 9: 1066-1079.
15

Th e study examines the eff ects of soil drainage on 

carbon accumulation and allocation across a Picea 

mariana fi re-infl uenced chronosequence in Manitoba, 

Canada. Two stand conditions (dry and wet) are 

compared across seven diff erent stands. Stands 

originated after stand-replacing wildfi res in 1850, 

1930, 1964, 1981, 1995, and 1998. Results show 

signifi cantly greater (p<.05) overstory biomass, MAI, 

woody debris mass, and litterfall for the dry stands; 

the wet stands have signifi cantly greater (p<.05) 

bryophyte, understory, and forest fl oor C pools. Total 

aboveground biomass for dry stands ranges from 

0.6-71.7 Mg C/ha (depending on age), but this is 

signifi cantly greater (p<.05) than the equivalent range 

for wet stands (0.3-31.2 Mg C/ha). Th is is within 

the range of other studies of boreal Picea mariana

stands (115-year-old P. mariana stands at BOREAS 

study sites have biomass C pools of 70.6 Mg C/

ha in northern dry areas, and 67.6 /17.3 Mg C/

ha in southern dry/wet areas, respectively). Biomass 

C pool estimates of P. mariana stands in western 

Canada range from 5.0-54.0 Mg C/ha; world P. 

mariana estimates range from 7.9-118.7 Mg C/ha. 

Total ecosystem C accumulation for the oldest stands 

(151 years old) is 533.5 Mg C/ha for dry stands and 

470.4 Mg C/ha for wet stands. Th e importance of soil 

drainage and age structure on temporal and spatial C 

distribution in these stands must be taken into account 

in large-scale C budget modeling.

II. Models

39. Akselsson, C.; Westling, O.; Sverdrup, H.; 

Gundersen, P. 2007. Nutrient and carbon 

budgets in forest soils as decision support in 

sustainable forest management. Forest Ecology 

and Management. 238: 167-174.

Th is study compares the potential for climate change 

mitigation in Swedish forests from C sequestration 

in forest ecosystems vs. fossil fuel substitution from 

bioenergy derived from whole-tree harvesting. Authors 

argue that sequestration in forest ecosystems is a 

method of delaying emissions, and therefore not a 

sustainable solution. Results indicate that biofuel from 

whole-tree harvesting can substitute for 3.6 million 

metric tons of oil/yr, which equates to emissions 

savings of 11 million tons CO2e/yr. Sequestration in 

forest soils currently yields a sink of 100 kg C/ha/yr, or 

2 million tons C/yr (7.34 million tons CO2e), but the 

study warns that this sink is not sustainable, and longterm capacity is not substantial. Whole-tree harvesting 

would lead to net losses in nutrients (specifi cally N and 

base cations), and compensatory fertilization would be 

required.

40. Alig, R.J.; Adams, D.M.; McCarl, B.A. 2002. 

Projecting impacts of global climate change on 

the U.S. forest and agriculture sectors and carbon 

budgets. Forest Ecology and Management. 169: 

3-14.

Th is study uses the FASOM model and fi nds that 

impacts to the United States forest sector by climate 

change are relatively small. Results agree with those 

of other studies, suggesting that currently in the 

United States, existing variation and yields exceed 

variations caused by climate change. Th e model 

assumes an inelastic demand for wood products 

and predicts changes in United States forest carbon 

inventory ranging from +0.1 to -0.2 percent through 

2020, and -1.0 percent to -1.7 percent through 2050, 

due to higher harvesting levels. Th e model does not 

consider climate feedback and changes to disturbances 

and assumes that timber production will “migrate” 

from north to south according to climate-induced 

productivity changes.

41. Backéus, S.; Wikström, P.; Lämås, T. 2005. 

A model for regional analysis of carbon 

sequestration and timber production. Forest 

Ecology and Management. 216: 28-40.

Th is study presents a local/regional optimization 

model (created using linear program formulation 

that is considered a mixture of model 1 and model 

2 formulation for linear modeling) that incorporates 

economic values for timber production and C 

sequestration (but does not incorporate eff ects of 

climate change). Th e model was developed using 

individual tree- and stand-level data from the 

Swedish National Forest Inventory and uses linear 

programming to develop optimal stand management 

solutions based on C prices. Harvesting generally 
16

decreases and landscape C storage increases as C prices 

increase; total storage for 3.2 Mha of boreal forest in 

Sweden ranges from 1.48-2.05 Mt C/yr.

42. Baral, A.; Guha, G.S. 2004. Trees for carbon 

sequestration or fossil fuel substitution: the issue 

of cost vs. carbon benefi t. Biomass and Bioenergy. 

27: 41-55.

Th is study compares the potential for mitigation 

through aff orestation without harvest vis-à-vis 

bioenergy using growth conditions and species typical 

of the southeastern United States. Diff erences over a 

100-year period are signifi cant: biomass-based BIG/

STIG (biomass-integrated gasifi ed/steam-injected 

gas turbine substituting) technology, if commercially 

available, cost-eff ective, and substituted for coal-fi red 

steam-electric power, could result in an additional C 

benefi t of 3.03 Mg C/ha over traditional aff orestation 

without harvest. Th is study notes that fi ndings should 

be considered theoretical until technology is proven as 

commercially viable and cost eff ective.

43. Bateman, I.J.; Lovett, A.A. 2000. Estimating and 

valuing the carbon sequestered in softwood and 

hardwood trees, timber products and forest soils 

in Wales. Journal of Environmental Management. 

60: 301-323.

Th is study stresses the importance of full C accounting 

in creating scientifi cally sound policy and states that C 

in aboveground biomass, wood products and waste, and 

soils must be considered in any study of C fl ux. Th e 

model demonstrates that economic drivers, specifi cally 

in the form of discount rates, exert the most infl uence 

on total C storage and emission values (lower discount 

rates increase fl ux). Other factors that infl uence C 

fl ux, in descending order of importance, are forest 

management, timber growth rates, and species 

selection. Planting on peat soils can stimulate large C 

releases that outweigh C sequestered in live wood.

44. Battaglia, M.; Sands, P.; White, D.; Mummery, 

D. 2004. CABALA: a linked carbon, water 

and nitrogen model of forest growth for 

silvicultural decision support. Forest Ecology and 

Management. 193: 251-282.

Th is paper develops and validates a robust, generally 

applicable, and accurate silvicultural decision-support 

system (model) that links C, water, and N fl ows 

through the atmosphere, biomass, and soils. A key 

diff erence between CABALA and other models is that 

it links aboveground and belowground processes. Th e 

process functions included in CABALA also provide a 

framework for localizing the model, which is generally 

suitable for decadal-scale studies, but perhaps not for 

long-term sustainability studies.

45. Bergh, J.; Freeman, M.; Sigurdsson, B.; Kellomäki, 

S.; Laitinen, K.; Niinistö, S.; Peltola, H.; Linder, S. 

2003. Modeling the short-term eff ects of climate 

change on the productivity of selected tree 

species in Nordic countries. Forest Ecology and 

Management. 183: 327-340.

Low primary productivity is characteristic of boreal 

forests, but increasing CO2 concentrations and 

temperatures associated with climate change could 

infl uence productivity and future boreal C balance. 

Th e study reports on simulations on coniferous (Pinus 

sylvestris, Picea abies) and deciduous broadleaf (Fagus 

sylvatica, Populus trichocarpa), stands across a climatic 

gradient in northern Europe. Scenarios modeled 

were control (0 °C and 350 ppm), +2 °C and 350 

ppm, +4 °C and 350 ppm, 700 ppm, +2 °C and 700 

ppm, +4 °C and 700 ppm. Coniferous species are 

responsive to increased temperatures, which induced 

an earlier start to photosynthetic capacity recovery 

and increased the length of spring recovery. Th e +2 

°C and +4 °C scenarios increased respiration rates 

throughout the year for all sites and species, but also 

generally increased water demands and water defi cit, 

which in turn, reduced NPP in some scenarios. 

NPP of P. sylvestris and P. abies increased by 24-37 

percent in the +4 °C scenario. Elevated CO2 scenarios 

increased NPP for coniferous species on some sites 

by 25-40 percent and increased NPP of Denmark 

beech sites by 58 percent, primarily by improving 

water use effi ciency. Th is paper stresses the need for 

research on low-temperature eff ects and soil-nutrient 

feedback mechanisms (increased growth naturally leads 

to increased nutrient demand) to avoid unrealistic 

predictions of C gains or climate change eff ects in 

boreal forest ecosystems.
17

46. Bond-Lamberty, B.; Peckham, S.D.; Ahl, D.E.; 

Gower, S.T. 2007. Fire as the dominant driver of 

central Canadian boreal forest carbon balance. 

Nature. 450: 89-92.

Th is study employed the Biome-BGC process model 

to study the eff ects of changing environmental 

conditions (changing climate, increased CO2

concentrations, fi re disturbances) on NPP, net biome 

production, vegetation competition, and C balance 

on 100 Mha of Canadian boreal forest dominated 

by Picea mariana and Populus tremuloides. Fire 

disturbance has increased concentrations of deciduous 

trees at the expense of coniferous trees, and that the 

C balance of the region from 1948-2005 has been 

driven primarily by fi re disturbance. Th e authors 

conclude that climate-related changes have been 

driven primarily by increased fi re frequency, which has 

aff ected the variability but not the mean of the region’s 

C balance over time, and that the region has not yet 

experienced direct ecophysiological changes driven by 

climate change.

47. Böttcher, H.; Freibauer, A.; Obersteiner, M.; 

Schulze, E.-D. 2008. Uncertainty analysis of 

climate change mitigation options in the forestry 

sector using a generic carbon budget model. 

Ecological Modelling. 213: 45-62.

Th is study presents a generic dynamic forestry model 

(FORMICA) that can be used to model forest C 

dynamics and sink potential of landscape forest C 

pools, as well as from harvested wood products and 

fossil fuel substitution.

48. Brack, C.L. 2002. Pollution mitigation and 

carbon sequestration by an urban forest. 

Environmental Pollution. 116: 195-200.

Th is paper delivers a case study that enumerates and 

economically quantifi es the benefi ts of an urban 

forest to the southeast Australian city of Canberra. 

Th e study uses a model to predict C sequestration by 

400,000 urban trees and converts that fi gure into a 

Kyoto-commitment-driven dollar value over 4 years 

(U.S. $300,000) and demonstrates that the majority 

of the value of urban trees derives from energy savings, 

pollution amelioration, and hydrology amelioration 

(estimated at U.S. $20-67 million for the city of 

Canberra from 2008-2012).

49. Bravo, F.; Bravo-Oviedo, A.; Diaz-Balteiro, 

L. 2008. Carbon sequestration in Spanish 

Mediterranean forests under two management 

alternatives: a modeling approach. European 

Journal of Forest Research. 127: 225-234.

Th is study examines C sequestration and associated 

economic impact on two diff erent even-aged 

management alternatives (long and short rotation) 

on two simulated stands. One stand was composed 

of Pinus sylvestris and the other of Pinus pinaster; 

both include numerous site quality indexes. In each 

case, the study simulates a clear cut followed by 

natural regeneration, precommercial thinning, and 

a low thinning regimen to achieve stand densities 

of 1,600 trees/ha in P. sylvestris and to 1,500 trees/

ha in P. pinaster. Increasing rotation length on poor 

quality sites allows trees to store as much C as shorter 

rotations on high quality sites, and combined longer 

rotations and gains in C revenue can make poor 

quality sites profi table while creating a positive impact 

on biodiversity. In all circumstances, increasing 

rotation length increases the proportion of C stock in 

the fi nal harvest relative to total fi xed C.

50. Briceño-Elizondo, E.; Garcia-Gonzalo, J.; Peltola, 

H.; Kellomäki, S. 2006. Carbon stocks and 

timber yield in two boreal forest ecosystems 

under current and changing climatic conditions 

subjected to varying management regimes.

Environmental Science & Policy. 9: 237-252.

Th e study uses a process-based ecosystem model 

(FINNFOR) to determine the impact of thinning 

regimes and changing climate on forest ecosystem 

C stocks. Simulations were performed over a 100-

year period for Pinus sylvestris, Picea abies, and Betula 

pendula in Finnish boreal forests. Across each of three 

climate scenarios (one current and two projection 

models: HadCM2 and ECHAM4), all three species 

demonstrated maximum C sequestration when 

traditional thinning regimens were most drastically 

altered to allow for a 30 percent increase in forest 

stock. Th e fi ndings demonstrate and validate the 
18

hypothesis that C stocks in forest ecosystems can 

be increased by applying a proper thinning regimen 

without increasing rotation length or experiencing 

losses in timber production. Th is study recommends a 

“no regrets” option of increasing thinning thresholds, 

thereby increasing total stocks and carbon sequestered.

51. Briceño-Elizondo, E.; Garcia-Gonzalo, J.; Peltola, 

H.; Matala, J.; Kellomäki, S. 2006. Sensitivity of 

growth of Scots pine, Norway spruce and silver 

birch to climate change and forest management 

in boreal conditions. Forest Ecology and 

Management. 232: 152-167.

Authors use a process-based ecosystem model 

(FINNFOR) linking C, N, energy, and water fl ows 

in trees and soils to examine climate change eff ects on 

tree growth and timber production for three species in 

northern and southern Finnish boreal forests. Th inning 

regimes that were less intensive than the traditional 

thinning level under two projected climate change 

scenarios resulted in universal increases in tree growth 

and timber yield compared to a baseline climate 

scenario. Gains generally were larger in the north, 

where forests benefi t more from warmer temperatures.

52. Briceño-Elizondo, E.; Jäger, D.; Lexer, M.J.; 

Garcia-Gonzalo, J.; Peltola, H.; Kellomäki, S. 2008. 

Multi-criteria evaluation of multi-purpose stand 

treatment programmes for Finnish boreal forests 

under changing climate. Ecological Indicators. 8: 

26-45.

Th is study uses MCA within the FINNFOR model to 

demonstrate that higher stocking levels, in the form 

of unthinned stands or maximally increased thinning 

thresholds (30 percent increase in upper and lower 

limits of basal area), result in greater utility compared 

to a BT treatment program; BT (0,0) is the standard/

recommended guideline for thinning in Finland. Th e 

study emphasizes the importance of reconsidering 

current forest management techniques (specifi cally 

thinning regimes) in light of changing climate 

scenarios, to maximize C storage and timber yield.

53. Chen, W.; Chen, J.M.; Price, D.T.; Cihlar, J.; 

Liu, J. 2000. Carbon off set potentials of four 

alternative forest management strategies in 

Canada: A simulation study. Mitigation and 

Adaptation Strategies for Global Change. 5: 143-

169.

Th is study uses an integrated terrestrial ecosystem 

C budget model (InTEC) to assess the mitigation 

potential of four alternative forest management 

scenarios in Canada. In order of greatest benefi t: 

amendment application (5 kg N/ha/yr) on 125m ha 

of semi-mature boreal forests (58 Tg C/yr); promptly 

reforesting disturbed land (57 Tg C/yr); increasing 

harvest levels by 20 percent to create bioenergy to 

off set fossil fuel use (11 Tg C/yr); and aff oresting all 

7.2 million ha of marginal agricultural land (8 Tg 

C/yr). Th is study presents experimental evidence to 

support theoretical gains from N application, though 

other studies show that the response of forests to 

fertilization remains highly variable.

54. Chladná, Z. 2007. Determination of optimal

rotation period under stochastic wood and 

carbon prices. Forest Policy and Economics. 9: 

1031-1045.

Th is publication presents a model to optimize forest 

management. Th e study demonstrates that optimal 

rotation length is closely connected to timber and C 

prices and price behavior, discount rate, and whether 

the forest landowner is responsible for C emissions at 

harvest.

55. Creedy, J.; Wurzbacher, A.D. 2001. Th e 

economic value of a forested catchment with 

timber, water and carbon sequestration benefi ts. 

Ecological Economics. 38: 71-83.

Th is study develops a mathematical model to 

optimize the net present value of timber, water, and 

aboveground sequestration benefi ts for a forested 

catchment in Victoria, Australia. Allowing for various 

carbon price and discount rate inputs, the model 

demonstrates that accounting for nontimber benefi ts 

can signifi cantly lengthen optimal rotation length.

56. De Jong, B.H.; Masera, O.; Olguín, M.; 

Martínez, R. 2007. Greenhouse gas mitigation 

potential of combining forest management and 
19

bioenergy substitution: A case study from Central 

Highlands of Michoacan, Mexico. Forest Ecology 

and Management. 242: 398-411.

Th e study employs the CO2FIX v 3.1 modeling 

software to determine the optimal strategy for 11,000 

ha of native predominantly Pinus-Quercus forests in 

central Mexico. Th e combination “oak conservation 

+ bioenergy” strategy provides the most C gain 21.6- 

42.9 tons C/ha. Th e oak conservation strategy stops 

yielding C gains (8.2 -19.3 tons C/ha) after about 40 

years, while the combination strategy yields gains of 

1.36 tons C/ha/yr continuously.

57. De Jong, B.H.J.; Tipper, R.; Montoya-Gómez, G. 

2000. An economic analysis of the potential for 

carbon sequestration by forests: evidence from 

southern Mexico. Ecological Economics. 33: 313-

327.

Th is study uses economic modeling to determine the 

potential for enhanced mitigation through improved 

forest management and other techniques on 600,000 

ha in southern Mexico. An additional 63 Mg C/

ha can be sequestered at a cost of U.S. $15/ton C 

($15/Mg C). Roughly 80 percent of these gains are 

directly related to improved forest management. Th is 

study also emphasizes the importance of a baseline in 

determining extra C stored and the cost-eff ectiveness 

of forest-based C storage.

58. Dean, C.; Roxburgh, S.; Mackey, B.G. 2004. 

Forecasting landscape-level carbon sequestration 

using gridded, spatially adjusted tree growth.

Forest Ecology and Management. 194: 109-129.

Th is publication reports on a model (CAR4D) to scale 

up C sequestration forecasts to the landscape level 

while accounting for nonlinearities, environmental 

variability, and temporal and spatial scaling issues that 

often lead to inaccuracies in landscape-level forecasts. 

C storage in Eucalyptus regnans stands peaks about 130 

years after establishment, at which point the forest 

becomes a net source. A C bank can be maintained at 

a stable level by mixing old and young stands across a 

landscape. A wider profi le of forest stand ages results 

in a more stable C bank forecast. Results also indicate 

when and how models can be improved through 

additional collection of fi eld data.

59. Díaz-Balteiro, L.; Romero, C. 2003. Forest 

management optimization models when carbon 

captured is considered: a goal programming 

approach. Forest Ecology and Management. 174: 

447-457.

Th is study presents a model that incorporates 

C sequestration objectives into optimal forest 

management and highlights some inherent tradeoff s, 

including unacceptable results that derive from a 

singular focus on optimizing any one criterion. It 

includes an optimization case study from a Pinus 

sylvestris dominated forest in north-central Spain.

60. Environmental Protection Agency. 2005. 

Greenhouse gas mitigation potential in U.S. 

forestry and agriculture. Washington, DC: U.S. 

Environmental Protection Agency. 154 p.

Although this study employs a comprehensive and 

dynamic model (Forest and Agriculture Sector 

Optimization Model with Greenhouse Gases—

FASOMGHG) and thus warrants categorization in 

this section, the study could easily fi t into section 4 

(Literature reviews and meta-analyses) because of its 

comprehensive, meta-analytical nature. Th e study 

integrates biological and economic data to demonstrate 

that the United States forestry and agricultural sectors 

together constitute a major C sink (~830 Tg C/

yr), over 90 percent of which is on forest land. Sink 

strength on forest land compensates for and outweighs 

releases due to wildfi res, harvests, and conversions 

to nonforest use. Economic modeling demonstrates 

the possibility for increased terrestrial storage given 

adequate GHG prices and certain assumptions 

about landowner behavior. Th e study suggests that 

at lower price ranges (less than or equal to $5/

tonCO2e), agricultural soil sequestration and forest 

management for increased C storage are “dominant 

mitigation strategies” and that aff orestation becomes 

the dominant strategy at higher (>$15/tonCO2e) 

prices. Data from the past several years generated by 

the Chicago Climate Exchange and over-the-counter 

carbon markets show that $5/tonCO2e has barely 
20

been suffi cient to cover costs, stimulate interest, and 

dissipate owner reluctance/uncertainty about engaging 

in markets and/or encumbering his or her land with 

long-term commitments.

61. Eriksson, E.; Gillespie, A.R.; Gustavsson, L.; 

Langvall, O.; Olsson, M.; Sathre, R.; Stendahl, 

J. 2007. Integrated carbon analysis of forest 

management practices and wood substitution. 

Canadian Journal of Forest Research. 37: 671-681.

Th is study examines the eff ects of three forest 

management scenarios (traditional, intensive 

management, and intensive fertilization), three 

slash-management practices (no removal, removal, 

and removal with stumps), two forest-product uses 

(construction material and biofuel), and two reference 

fossil fuels (coal and natural gas) on the total C 

balance of a Picea abies forest. Th is study combines a 

forest-growth, forest-soil, and product-use model to 

represent the system, and fi nds that forest-product 

usage has much greater impact on the total carbon 

budget than changes in accumulation in biomass and 

soils due to various types of management. Th e greatest 

overall C gains are achieved through a combination 

strategy of stand fertilization, harvest with slash and 

stumps removed, and timber converted into long-term 

construction wood products and then substituted 

for coal-based fuel after its useful life as construction 

materials. Th is strategy generated a savings of more 

than 3 Mg C/ha/yr above a baseline (excludes 

fertilization or stump removal, and substitutes wood 

products for natural gas). Results support the idea that 

intensively managed forests can signifi cantly mitigate 

climate change and that most of the benefi t derives 

from substituting wood for more C-intensive products.

62. Felzer, B; Kicklighter, D.; Melillo, J.; Wang, C.; 

Zhuang, Q.; Prinn, R. 2004. Eff ects of ozone on 

net primary production and carbon sequestration 

in the conterminous United States using a 

biogeochemistry model. Tellus B. 56: 230-248.

Th is study evaluates the eff ects of ozone on NPP and 

terrestrial C sequestration in the coterminous United 

States using the Terrestrial Ecosystem Model. It 

estimates a 2.6-6.8 percent mean reduction in annual 

NPP from ozone which translated into a reduction in 

United States forest C sequestration of 18-38 Tg C/yr 

since the 1950s.

63. Fujikake, I. 2007. Selection of tree species for 

plantations in Japan. Forest Policy and Economics. 

9: 811-821.

Th is study examines the eff ect of economic factors 

on decisions about which species to plant in Japanese 

plantation forests. Changes in stumpage prices and 

forestry wages have been the primary drivers in 

a switch from Sugi to Hinoki. Hinoki forests are 

slower growing, generally allow for less understory 

development, and have faster litter decomposition and 

greater soil erosion rates, making them less robust C 

sinks. If forest C and other ecosystem benefi ts were 

quantifi ed and valued economically, managers of 

plantation forests could make long term decisions that 

are more ecologically sound.

64. Fürstenau, C.; Badeck, F.; Lasch, P.; Lexer, 

M.; Lindner, M.; Mohr, P.; Suckow, F. 2007. 

Multiple-use forest management in consideration 

of climate change and the interests of stakeholder 

groups. European Journal of Forest Research. 126: 

225-239.

Th e MCA method is paired with the 4C (Forest 

Ecosystems in a Changing Environment) growth 

model to determine the utility of a range of forest 

management scenarios under a variety of climate 

conditions to three separate stakeholder groups 

(private forest owner, public forest manager, and 

environmental organization). Th e greatest overall 

increase in total C sequestration was greatest for 

the “no management/conservation strategy”, even 

when C in soil and long-term wood products from 

other scenarios was considered. However, the utility 

of this management scenario was low for all but 

the environmental organization, and this one-time 

increase in C storage was vulnerable to release through 

climate-related or other disturbances. Also, the MCA 

method is designed to maximize the utility of adaptive 

management decisions to various stakeholder groups in 

an era of uncertain climate changes, so C sequestration 

is only one component of a number of functions (e.g., 
21

timber yield, groundwater recharge, and biodiversity 

factors) that must be considered. Outcomes depend 

entirely on specifi c circumstances and local conditions.

65. Garcia-Quijano, J.F.; Deckmyn, G.; Ceulemans, 

R.; Van Orshoven, J.; Muys, B. 2008. Scaling 

from stand to landscape scale of climate 

change mitigation by aff orestation and forest 

management: A modeling approach. Climatic 

Change. 86: 397-424.

A process-based model (SECRETS) designed to 

overcome stand-to-landscape scaling problems is 

linked with a C accounting model (GORCAM) to 

fully consider the carbon implications and eff ects 

of forest management, aff orestation, and bioenergy 

production. Th e model accurately predicts C stocks in 

various pools for current environmental conditions, 

but comparison with fi eld data suggests that accurate 

predictions under changing climate scenarios require 

greater functionality. Results suggest that under 

current climate conditions, the best scenario for 

both sequestering and avoiding emissions involves a 

combination of conservation of old-growth reserves, 

use of existing multifunctional forest, and “short 

rotation coppice” for bioenergy production. Soil C and 

litter pools present a modeling challenge and related 

inaccuracies in these pools may amplify uncertainties.

66. Garcia-Gonzalo, J.; Peltola, H.; Zubizarreta 

Gerendiain, A.; Kellomäki, S. 2007. Impacts of 

forest landscape structure and management 

on timber production and carbon stocks in the 

boreal forest ecosystem under changing climate. 

Forest Ecology and Management. 241: 243-257.

Th e study uses a process-based model (FINNFOR) to 

determine the C sequestration sensitivity of a boreal 

forest ecosystem to diff erent age class distributions and 

thinning regimes. Th e eff ects of the HadCM2 climate 

scenario (increased temperature, precipitation, and 

atmospheric CO2 concentration) increased de facto

forest productivity (total aboveground stock increased 

while soil C decreased due to higher respiration, but 

the overall eff ect was a net gain in ecosystem C). 

Unthinned stands result in maximum ecosystem 

sequestration. Although thinning resulted in ecosystem 

C losses, a shift in thinning from BT (0,0) to BT (30, 

30), or a 30 percent increase in the thinning threshold 

that allowed for higher stocking densities, resulted in 

notable C benefi ts (13 Mg C/ha) over the 100-year 

simulation period under both current and future 

climate scenarios. Initial age class distribution generally 

did not result in substantial diff erences in forest C 

stock over 100 years.

67. Gutrich, J.; Howarth, R.B. 2007. Carbon 

sequestration and the optimal management 

of New Hampshire timber stands. Ecological 

Economics. 62: 441-450.

A model is used to examine decisions related to 

harvesting/rotation length New Hampshire in light 

of the economic tradeoff s between timber and C 

sequestration values. Results show that optimal 

rotation length is extended from 16 to 133 years 

depending on forest type when forest C storage is 

valued between $25-75 per ton C, respectively. As a 

second scenario, the study cites the UNFCCC goal 

to stabilize atmospheric CO2 concentrations at a level 

that would avoid catastrophic harm. At 580 ppm, 

Howarth (2001) estimates C prices to be $227-570/ 

metric ton C; at such prices, the study estimates that 

optimal rotation length of stands in New Hampshire 

would range from 222-391 years. Partial harvesting 

rather than clearcut-harvesting provides “high net 

benefi ts under a variety of circumstances.” Th e model 

only incorporates optimal management at the stand 

level, and does not consider regional-level impacts 

(leakage, etc.); model valuation of timber could be 

improved and may not represent conditions on a 

specifi c timber stand.

68. Harmon, M.; Marks, P. 2002. Eff ects of

silvicultural practices on carbon stores in 

Douglas-fi r-western hemlock forests in 

the Pacifi c Northwest, USA: results from a 

simulation model. Canadian Journal of Forest 

Research. 32: 863-877.

Th is study uses the model STANDCARB to simulate 

the eff ects of fi ve treatments on C stores in stands 

of Pseudotsuga menziessii and Tsuga heterophylla in 

Oregon, and found that three factors in particular 
22

(detritus removed by slash burning, amount of mass 

harvested, and rotation length, listed in order of 

decreasing importance) greatly aff ect landscape carbon 

stores. Th ere was a notable decline in landscape C 

following conversion/disturbance/management of 

old-growth forests. Agricultural land stored only 15 

percent of total C capacity whereas “forests protected 

from fi re” stored 93 percent of total capacity. 

Th erefore, converting agricultural land to forest 

increased landscape C stores.

69. Hennigar, C.R.; MacLean, D.A.; Amos-Binks, L.J. 

2008. A novel approach to optimize management 

strategies for carbon stored in both forests and 

wood products. Forest Ecology and Management. 

256: 786-797.

An analysis is presented which demonstrates the 

importance of full C accounting when attempting 

to optimize mitigation. It evaluates four separate 

objectives for a modeled forest in New Brunswick, 

Canada, (using Remsoft Spatial Planning System) and 

fi nds that a scenario to maximize carbon in forest + 

products + substitution resulted in storage of 0.228 

Mg C/ha/yr; a scenario to maximize harvest resulted in 

storage rates of 0.126 Mg C/ha/yr.

70. Huang, C.-H.; Kronrad, G.D. 2001. Th e cost 

of sequestering carbon on private forest lands.

Forest Policy and Economics. 2: 133-142.

Th is study uses the forest stand simulator model 

PTAEDA2 to analyze the costs of C sequestration of 

Pinus taeda, comparing the SEV of biologically vs. 

economically optimal rotation lengths. Th e former 

maximizes C stored on the forest and in long-lived 

wood products; the latter maximizes the economic 

value of sawtimber. Th e objective was to determine 

the maximum mean annual increment of these two 

goals. Results show a diff erence of as much as 0.79 

Mg C/ha. Depending on site quality class (50, 60, 

70, 80, 90) and the discount rate used, the price per 

additional ton of sequestered C on forest land ranged 

from $0.84 to $72.79/ha. Th is is the price that one 

would need to pay forest managers to incentivize them 

to switch from economically to C-optimal rotation 

lengths). Th e sequestration potential and associated 

price for incentivizing landowners to restock barren 

land was 1.03-3.77 Mg C/ha and $0.74 - $27.32/ha, 

respectively. Th e cost of sequestering an additional ton 

of C intensively managed land was $4.18 - $181.27/

ha. Findings assumed an initial age of 25 years, 

maximum rotation of 60 years, 0-3 thinnings (fi rst 

thinning would be row and from below; subsequent 

thinnings, if any, would be from below) and thinning 

intensities of 20, 25, 30, or 35 percent of basal area 

removed.

71. Hurteau, M.D.; Koch, G.W.; Hungate, B.A. 2008. 

Carbon protection and fi re risk reduction: toward 

a full accounting of forest carbon off sets. Frontiers 

in Ecology and the Environment. 6: 493-498.

Th is study quantifi es the CO2 released from the four 

largest U.S. wildfi res in 2002 (Rodeo–Chediski, 

Hayman, Biscuit, and McNally) in light of forest 

management practices that might mitigate or reduce 

the likelihood of such large releases. Th ese fi res charred 

508,000 ha, 92,000 of which were burned severely. 

Previous studies and models were used to estimate C 

storage for unmanaged forests typical of those that 

burned, and derive estimates of C storage per tree 

component and per hectare. Models and combustion 

factors were also used to estimate post-fi re C storage 

for burned thinned and unthinned stands. Results 

showed that 4.2-6.1 Mt CO2e were released in fi res 

from unthinned stands. Th inning these stands would 

have released 3.9 Mt CO2e but reduced live-tree fi re 

emissions by 0.07-0.3 Mt CO2e.

72. Hynynen, J.; Ahtikoski, A.; Siitonen, J.; Sievänen, 

R.; Liski, J. 2005. Applying the MOTTI simulator 

to analyze the eff ects of alternative management 

schedules on timber and non-timber production. 

Forest Ecology and Management. 207: 5-18.

Th is MOTTI stand model was used to simulate the 

eff ects of three stand management alternatives on a 

35-year old Picea abies-dominated stand in southern 

Finland. Management alternatives included a pure 

spruce stand with commercial thinning (x2) and 

harvest (at age 70) with the goal of maximizing wood 

production, a mixed stand with commercial thinning 

(x2) and harvest (at age 70) with a goal of enhancing 
23

biodiversity, and an unmanaged stand that was not 

treated. Over the 70-year simulation period, total soil 

C pool was roughly the same in pure and mixed stands 

and only slightly higher in no management stands. 

Th e tree C pool also was about the same in pure 

and mixed stands, but much higher in unmanaged 

stands (about 215 for pure/mixed vs. 270 Mg C/ha). 

Biodiversity was 50 and 88 percent higher in mixed 

and unmanaged stands, respectively. Th ese results 

based on underlying assumptions and thus subject to 

errors and uncertainties. Also, the biodiversity fi ndings 

may not be applicable to all boreal forest ecosystems.

73. Janisch, J.; Harmon, M. 2002. Successional 

changes in live and dead wood carbon stores: 

implications for net ecosystem productivity. Tree 

Physiology. 22:77-89.

Th is study uses data from a 500-year-old 

chronosequence in the Pacifi c Northwest to model the 

eff ects of forest management on the woody component 

of NEP, specifi cally C recovery horizons following 

management/disturbance. Transitioning from old 

growth to younger managed forest stands results in 

signifi cant C losses that are not recovered for many 

decades. Eighty-year-old regenerating stands stored 

half the C of 500-year old growth forests. Th e study 

notes that such scientifi c fi ndings must be carefully 

considered when designing mitigation projects to avoid 

adverse eff ects and net releases of C to the atmosphere.

74. Jayawickrama, K.J.S. 2001. Potential genetic

gains for carbon sequestration: A preliminary 

study on radiate pine plantations in New 

Zealand. Forest Ecology and Management. 152: 

313-322.

Th is study reports on modeled C gains for Pinus 

radiata plantations in New Zealand as a result of two 

diff erent levels of genetic improvement (selecting 

the best 50 of 500 parents, and the best 10 of 1,000 

parents). Th ese C gains were mapped over diff erent 

geographical regions and management strategies over 

a rotation length of 28 years (which included two 

thinnings and one pruning for all locations). Modeled 

long-term average gains are as much as 22 percent 

for the more stringent improvement (best 10 of 1000 

parents). Increases ranged from 19.0 to 29.3 Mg C/ha 

depending upon site and using a standard (direct saw 

log) management regime. For the same site, the 28-

year gains were highest when a plant-and-leave “lowcost” management approach was used.

75. Jiang, H.; Apps, M.J.; Peng, C.; Zhang, Y.; Liu, J. 

2002. Modelling the infl uence of harvesting on 

Chinese boreal forest carbon dynamics. Forest 

Ecology and Management. 169: 65-82.

Th is paper examines modeled eff ects of three 

harvest/disturbance scenarios (no biomass removal, 

conventional harvesting, whole-tree harvesting) over 

100-year rotations, and the same scenarios but with 

rotation lengths of 30, 50, 100, 200, or 400 years on 

Larix (dominant)-Betula-Pinus-Quercus boreal forests 

in China. Th e “no biomass removal” scenario resulted 

in C gains of 303 and 330 Mg C/ha over conventional 

and whole-tree harvesting, respectively, over 100 years. 

Chinese boreal forests generally are more productive 

and contain more biomass than North American or 

Russian boreal forests.

76. Kaipainen, T.; Liski, J.; Pusssinen, A.; 

Karjalainen, T. 2004. Managing carbon sinks by 

changing rotation length in European forests.

Environmental Science & Policy. 7: 205-219.

Th e study uses the CO2FIX model to determine the 

eff ects and magnitude of increased rotation lengths on 

European forest C pools to determine how to achieve 

the largest eligible C sink for Kyoto Protocol-related 

commitments. Increased rotation lengths result in 

slight decreases in soil C stock in some regions and 

generally decrease wood product pools (except in U.K. 

Picea sitchensis forests). Depending on forest and site 

characteristics, rotation lengths should be extended 

by 20 years on 0.3-5.1 Mha of land to achieve the 

aforementioned sink.

77. Karjalainen, T.; Pussinen, A.; Liski, J.; Nabuurs, 

G.-J.; Eggers, T.; Lapveteläinen, T.; Kaipainen, 

T. 2003. Scenario analysis of the impacts of 

forest management and climate change on the 

European forest sector carbon budget. Forest 

Policy and Economics. 5: 141-155.
24

Th is study uses scenario analysis to predict the eff ects 

of two forest management (business as usual, or BAU; 

and multifunctional management, which included 

increased fellings, but also aff orestation projects and an 

emphasis on “nature-oriented management”) and two 

climate scenarios (current climate and HadCM2) on 

the European forest C budget from 1990-2050. Th e 

study encompasses 27 countries and 128.5 Mha. Both 

management scenarios showed larger total ecosystem 

gains under HadCM2 than current climate. Because 

of current climate change eff ects, current climate 

scenarios (which were developed several years ago) 

likely can be dismissed as improbable. Th e diff erence 

between multifunctional and BAU management 

scenarios in a HadCM2 climate in 2050 was 215 Tg, 

or about 1.67 Mg/ha. Authors note that uncertainties 

still exist and could be minimized with more accurate 

biomass expansion factor and that when scenario 

analysis is used, more attention should be focused on 

relative than on absolute scenario diff erences.

78. Karjalainen, T.; Pussinen, A.; Liski, J.; Nabuurs, 

G.-J.; Erhard, M.; Eggers, T.; Sonntag, M.; 

Mohren, G.M.J. 2002. An approach towards an 

estimate of the impact of forest management and 

climate change on the European forest sector 

carbon budget: Germany as a case study. Forest 

Ecology and Management. 162: 87-103.

Process-based models can be used to assess regional 

climate change impacts, which then can be scaled up 

to regional/country/continental levels using national 

forest inventory data along with the European forest 

information scenario (EFISCEN) model to investigate 

the eff ects of diff erent forest management scenarios 

on C budgets in an era of climate change. Germany is 

used as a case study.

79. Köhl, M.; Stümer, W.; Kenter, B.; Riedel, 

T. 2008. Eff ect of the estimation of forest 

management and decay of dead woody material 

on the reliability of carbon stock and carbon 

stock changes–A simulation study. Forest Ecology 

and Management. 256: 229-236.

IPCC reporting guidelines off er three options for 

accounting for C stocks in dead wood. Th is study 

demonstrates that diff erent approaches to timber 

utilization and harvesting and assumed decay rates 

(constant or accelerating) of dead wood greatly aff ect 

the amount of C released. When these eff ects were 

modeled for a given set of data from Austria, changes 

in C stock in dead woody material ranged from 3.1-4.4 

Mg C/ha/yr.

80. Kraxner, F.; Nilsson, S.; Obersteiner, M. 2003. 

Negative emissions from BioEnergy Use, Carbon 

capture and Sequestration (BECS)—the case 

of biomass production by sustainable forest 

management from semi-natural temperate 

forests. Biomass and Bioenergy. 24: 285-296.

Th e study proposes a negative emissions energy system 

using “semi-natural” temperate forests. It states that 

“Biomass Energy CO2 removal through long-term 

Sequestration” (BECS) provides an insatiable sink for 

C, thereby reducing atmospheric CO2 concentrations 

and risks from nonlinear climate changes. Stem 

biomass is used to generate energy, while other 

biomass is left in the forest to preserve nutrient 

balances and ecosystem integrity. C can be scrubbed 

and stored during the energy production process; 

a range of diff erent technology and carbon storage 

options is cited. C sequestration trajectories in unevenaged mixed beech-spruce stands are projected under 

four scenarios: with and without BECS, using various 

forest management regimes. Th inning and one-time 

planting with BECS generated the highest average 

annual sequestration of 2.5 Mg C/ha/yr. Th inning 

and one-time planting without BECS generated the 

lowest (0.85 Mg C/ha/yr) average, even lower than 

the control. Using BECS technology might produce a 

comparative insatiable and permanent sink of 1.65 Mg 

C/ha/yr, but the study cites the need for governmental 

and societal support.

81. Krcmar, E.; Stennes, B.; van Kooten, G.C.; 

Vertinsky, I. 2001. Carbon sequestration and land 

management under uncertainty. European Journal 

of Operational Research. 135: 616-629.

Th e study models potential C uptake on reforested 

and aff orested land, taking into account inherent 

uncertainties related to forest management. 
25

Uncertainties include: timber yield quantities related 

to silvicultural practices, climate change eff ects, and 

weather; imprecise C sink data; and “vague and 

changing societal values and preferences,” especially 

with respect to timber harvest and C sequestration 

levels. Modeled results indicate that parameters and 

assumptions signifi cantly impact land- use strategies 

and outcomes. Th e study concludes by stating that 

models are useful but cannot determine the best 

strategy, so the onus of determining inputs and 

assumptions and determining thresholds for risk lies 

with decision-makers.

82. Krcmar, E.; van Kooten, G.C.; Vertinsky, I. 

2005. Managing forest and marginal agricultural 

land for multiple tradeoff s: compromising 

on economic, carbon and structural diversity 

objectives. Ecological Modelling. 185: 451-468.

Th e study uses “compromise programming” to 

optimize net present value, nominal and discounted 

C sequestration totals, and biodiversity structure 

on a 70/30 percent coniferous/deciduous mixed 

boreal forest in British Columbia, Canada. Th e study 

experiments with balancing or maximizing all or 

certain of the four objectives and fi nd that signifi cant 

tradeoff s exist among achieving maximal sequestration, 

economic benefi t, and landscape structural diversity. 

Namely, maximizing short-term C uptake results in 

signifi cant compromises in diversity; maintaining a 

diverse landscape reduces potential economic and 

C sequestration. Land managers with diff erent risk 

tolerance levels will utilize diff erent land-use and 

management strategies.

83. Kurz, W.A.; Dymond, C.C.; Stinson, G.; 

Rampley, G.J.; Neilson, E.T.; Carroll, A.L.; Ebata, 

T.; Safranyik, L. 2008. Mountain pine beetle and 

forest carbon feedback to climate change. Nature. 

452: 987-990.

Th e study quantifi es the C impacts of a mountain pine 

beetle outbreak in British Columbia, Canada. Th e 

most recent beetle outbreak is an order of magnitude 

greater than other outbreaks due to an increased area 

of mature pine stands and climate change related 

favorable beetle conditions, and has resulted in 

massive pine mortality and commercial timber losses 

estimated at 435 million m3. Th is CBM-CFS3 model 

was used to estimate the combined eff ects of the beetle 

outbreak, forest fi res, and harvesting on south-central 

British Columbia’s forest C balance from 2000-2020 

(374,000 km2 of Pinus and Picea dominated forests). 

Th e cumulative eff ect caused a net release of 270 Mt, 

or 0.36 Mg C/ha/yr, thus turning these forests from a 

small sink to a substantial source. Th e study concludes 

that such climate-related disturbances may overwhelm 

northern forests’ ability to sequester and store 

atmospheric C, and that future large-scale modeling 

analyses must take this into account.

84. Kurz, W.A.; Stinson, G.; Rampley, G. 2008. 

Could increased boreal forest ecosystem 

productivity off set carbon losses from increased 

disturbances? Philosophical Transactions of the 

Royal Society B: Biological Sciences. 363: 2259-

2268.

Th is study uses the CBM-CFS3 model to compare the 

anticipated eff ects of climate change related increases 

in NPP vis-à-vis the impacts of increased fi re and other 

disturbances over a hypothetical 100,000 ha boreal 

forest landscape consisting of even-aged stands. Th e 

study examines two sets of 81 scenarios; each scenario 

set combines one of nine growth rate conditions with 

one of nine disturbance rate conditions. Th e fi rst set of 

scenarios features constant decomposition rates; in the 

second, decomposition rates increase proportionally 

with growth rates. Th e study models results to 200 

years, though focuses attention on results for the fi rst 

50 years. Th e fi rst set of scenarios suggests that climaterelated enhanced C sequestration can compensate 

for climate-related increased disturbances. However, 

the second scenario suggests that this will not occur 

should decomposition rates increase in proportion 

to increases in growth rate. In the fi rst scenario, a 

12 percent increase in growth rates is required to 

compensate for C losses associated with a 20 percent 

increase in annual burn area. However, the study 

concludes that projected increases in disturbance rates 

are likely to overwhelm growth enhancements such 

that “the boreal forest biome will act as a net source of 

C to the atmosphere if climate change brings about the 

projected increases in disturbance rates.”
26

85. Lasch, P.; Badeck, F.-W.; Suckow, F.; Lindner,M.; 

Mohr, P. 2005. Model-based analysis of 

management alternatives at stand and regional 

level in Brandenburg (Germany). Forest Ecology 

and Management. 207: 59-74.

Th e study tests and validates a physiologically based 

model (FORSEE—FOReST Ecosystems in a changing 

Environment, also known as 4C) parameterized for 

Fagus, Picea abies, Pinus sylvestris, Quercus, and Betula

using long-term experimental data from a forest in 

Germany. Th e model simulated the combined eff ects 

of growth, yield, management, and water on C budgets 

of managed forests over a 50-year horizon without fi nal 

harvest or stand regeneration under three management 

scenarios (without thinning but with removal of dead 

wood; with intensive thinning; and with moderate 

thinning) and two climate scenarios (current climate 

and ECHAM4 +1.4 °C change) to determine overall 

mitigative potential of the forest. Carbon stock 

assessed includes the sum of total biomass: total C in 

mineral soil and organic layer, harvested biomass, and 

deadwood. In both climate scenarios, the no-thinning 

management option results in greatest total C storage. 

Simulated climate change has little eff ect on total C 

storage (591.5 Tg C/ha total for current vs. 587.8 Tg 

C/ha total for climate change scenario). Under both 

climate scenarios, thinning results in less biomass in 

the forest, more in harvested wood products, and a 

reduction in soil carbon pools. Increased rotation 

lengths (160 yrs) results in greatest total carbon 

storage, but provides suboptimal timber revenues and 

quality. Projected increases in forest productivity due 

to longer growing seasons under a changing climate 

scenario are off set by signifi cantly reduced (-40 

percent) percolation rates. Th ere is limited potential 

for adaptive forest management in this region, which is 

characterized by dry conditions and poor site quality.

86. Law, B.E.; Th ornton, P.E.; Irvine, J.; Anthoni, 

P.M.; Van Tuyl, S. 2001. Carbon storage and 

fl uxes in ponderosa pine forests at diff erent 

developmental stages. Global Change Biology. 7: 

755-777.

Th e BIOME-BCG model was used to determine NPP, 

NEP, and C storage by component for two Pinus 

ponderosa stands in central Oregon: a 15-year-old stand 

in an old growth forest that had been clearcut and 

allowed to regenerate naturally and an unlogged stand 

of trees ranging from 50-250 years old. Aboveground 

tree C storage (bark, branch, stem, foliage) was 5.79 

Mg C/ha for the young stand and 107.91 Mg C/ha for 

the old stand. Fifteen percent of C in the young stand 

was stored as aboveground live biomass vs. 61 percent 

for the old stand. Th e old stand sequesters anywhere 

from 0.2-1.0 Mg C/ha/yr.

87. Law, B.E.; Sun, O.J.; Campbell, J.; Van Tuyl, S.; 

Th omton, P.E. 2003. Changes in carbon storage 

and fl uxes in a chronosequence of ponderosa

pine. Global Change Biology. 9: 510–524.

Th e BIOME-BCG model was used to determine the 

NPP, NEP, and C storage in aboveground and soil 

pools in Pinus ponderosa stands of four age classes 

(initiation, 9-23 years; young, 56-89 years; mature, 95-

106 years; and old, 190-316 years) in central Oregon. 

Total ecosystem C storage increases substantially—most 

of this increase is in aboveground biomass—until the 

stands are 150-200 years old, at which point the rate of 

storage slows but total storage continues to increase. Th e 

oldest age class (190-316 years) has a mean storage of 

176 Mg C/ha and 85 percent of biomass-based carbon 

storage is in older (100+ years) stands.

88. Law, B.E.; Turner, D.; Campbell. J.; Sun, O.J.; 

Van Tuyl, S.; Ritts, W.D.; Cohen, W.B. 2004.

Disturbance and climate eff ects on carbon stocks 

and fl uxes across Western Oregon USA. Global 

Change Biology. 10: 1429-1444.

Th is study employed both remote and fi eld-based 

observations with the BIOME-BCG model to estimate 

total C storage by Oregon forests, which was found 

to vary considerably by ecoregion and with respect 

to above/belowground partitioning. Total stock in 

Oregon is 2765 Tg C (337 Mg C/ha) and mean 

annual uptake is 13.8 Tg C/yr (1.68 Mg C/ha/yr).

89. Leighty, W.; Hamburg, S.; Caouette, J. 2006. 

Eff ects of management on carbon sequestration 

in forest biomass in Southeast Alaska. Ecosystems. 

9: 1051-1065.
27

Th e Tongass National Forest in Alaska is the largest 

area of old growth forest in the nation, containing 

8 percent of the C found in the forests of the 

coterminous United States and 0.25 percent of global 

forest ecosystem C. Th is study examines the eff ects of 

fi ve management scenarios (no harvesting; all lands 

harvested on a 100-yr rotation; all lands harvested 

on a 200-yr rotation; administratively available lands 

harvested on a 100-yr rotation; administratively 

available lands harvested on a 200-yr rotation) on total 

atmospheric C fl ux (all harvesting scenarios include 

gradual decay of harvested wood products, which were 

50/50 sawtimber and pulpwood) from 1995-2095. Th e 

model (spatially explicit combined GIS and FIA data 

model) was also run for a 200-year period from 1995-

2195. Results indicate that cessation of all harvesting 

on available lands results in additional sequestration 

of 0.03 – 0.25 Mg C/ha/yr. Th is fi ndings suggest that 

the Tongass could generate approximately as much 

revenue ($4-7million/yr) by eliminating harvesting and 

selling CERs for $20/Mg C (about $5.50 ton/ CO2e) 

as it does selling timber ($6.5million/yr in 2001). Th e 

policy questions yet to be resolved are: will and should 

National Forest System lands be eligible to generate C 

credits, and how will leakage and risk of reversals aff ect 

overall estimates of C benefi t?

90. Lenihan, J.M.; Bachelet, D.; Neilson, R.P.; 

Drapek, R. 2008. Simulated response of 

conterminous United States ecosystems to climate 

change at diff erent levels of fi re suppression, 

CO2 emission rate, and growth response to CO2. 

Global and Planetary Change. 64: 16-25.

Th is paper reports on a continental analysis of 

ecosystem response to simulated scenarios of various 

emission rates, CO2 fertilization eff ects, and levels 

of fi re suppression. Th ere are a variety of responses 

depending on potential inputs (including a variety 

of regional carbon gains and losses). Climate change 

eff ects remain uncertain and impacts and consequences 

depend on both ecosystem response and fi re 

management.

91. Liski, J.; Pussinen, A.; Pingoud, K.; Mäkipää, R.; 

Karjalainen, T. 2001. Which rotation length is 

favourable to carbon sequestration? Canadian 

Journal of Forest Research. 31: 2004-2013.

Rotation length can signifi cantly aff ect aboveground, 

soil, and wood product C pools. Th is study combines 

four models to determine the eff ects of simulated 

rotation length (60, 90, and 120 years over a 300-year 

horizon) on these three pools for Picea abies and Pinus 

sylvestris in a southern Finnish forest, while taking into 

account fossil C emissions and energy use in harvesting 

and product manufacturing. Results from the study 

indicate that for Pinus sylvestris, total sequestration is 

maximized (~ +2.5 percent) and emissions minimized 

by switching from a 90 (default) to 120-yr rotation 

length, but the tradeoff is a 10 percent decline in net 

revenues to landowners. Th e results for Picea abies

were less straightforward: whereas the shortest rotation 

(60 years) maximized the total carbon stock (in part 

because of signifi cant increases in soil C), it also 

resulted in the greatest harvest and manufacturingrelated emissions, which quickly negates ecosystem 

C stock surpluses. Increasing the rotation length to 

120 years results in slightly lower total C stocks (~ 

-1 percent), but substantially reduces harvest and 

manufacturing-related emissions. As a result, the latter 

strategy is recommended.

92. Luxmoore, R.J.; Hargrove, W.W.; Lynn Th arp, 

M.; Mac Post, W.; Berry, M.W.; Minser, K.S.; 

Cropper, W.P.; Johnson, D.W.; Zeide, B.; 

Amateis, R.L.; Burkhart, H.E.; Baldwin, V.C.; 

Peterson, K.D. 2002. Addressing multi-use 

issues in sustainable forest management with 

signal-transfer modeling. Forest Ecology and 

Management. 165: 295-304.

Th is publication describes a signal-transfer modeling 

approach developed for forests in the southeastern 

United States that provides a “means for gaining 

insight into possible large scale responses to impacts 

that have or can only be quantitatively investigated at 

a small scale.” Th is modeling approach can help land 

managers better determine management practices that 

will meet predefi ned sustainability goals related to 

water supply, C sequestration, timber production, and/

or biodiversity.
28

93. Luxmoore, R.J.; Th arp, M.L.; Post, W.M. 2008. 

Simulated biomass and soil carbon of loblolly 

pine and cottonwood plantations across a 

thermal gradient in southeastern United States.

Forest Ecology and Management. 254: 291-299.

A stand growth and soil C and N model (LINKAGES 

modifi ed to include RothC) are combined to simulate 

the eff ects of N fertilization and increased growingdegree-days on a 25-year Pinus taeda stand and for 

three consecutive 7-year rotations of Populus for 17 

sites in the southeastern United States. Results for both 

tree species showed marked increases in aboveground 

biomass, but C gains were not quantifi ed. Soil C stocks 

for both tree types were not aff ected by an increase in 

growing-degree-days. Th e authors suggest that “global 

warming may not initially aff ect soil C”, adding that 

more data are needed. Th e response of Populus to N 

fertilization was small but considered signifi cant: 12.9 

Mg C/ha to 13.7 Mg C/ha (equal to 0.04 Mg C/ha/yr 

over 21 years). Populus also showed signifi cant declines 

in relative response to N fertilization with increasing 

growing-degree-days, indicating that response generally 

is greater in cooler areas.

94. Magnani, F.; Consiglio, L.; Erhard, M.; Nolè, 

A.; Ripullone, F.; Borghetti, M. 2004. Growth 

patterns and carbon balance of Pinus radiate and 

Pseudotsuga menziesii plantations under climate 

change scenarios in Italy. Forest Ecology and 

Management. 202: 93-105.

Th e study validates a process-based growth model 

(HYDRALL) and simulates the eff ects of current 

and HadCM2 climate conditions on growth in 

three Pinus radiate and one Pseudotsuga menziesii

plantation in Italy. Th e climate change scenario 

increases forest growth, mean annual increment, and 

water-use effi ciency for all stands, and increases net 

ecosystem exchange only for the P. menziesii stand. C 

sequestration and/or storage is not reported.

95. Masera, O.R.; Cerón, A.D.; Ordóñez, A. 

2001. Forestry mitigation options for Mexico: 

Finding synergies between national sustainable 

development priorities and global concerns. 

Mitigation and Adaptation Strategies for Global 

Change. 6: 291-312.

Approximately 116 Mha of Mexico is covered by 

native forests; about 25.5 Mha is temperate forest land, 

66 Mha is semi-arid, and the remainder is tropical. 

Th is study simulates countrywide future emissions 

and forest-based sequestration using multiple model 

scenarios. Th e estimated “total mitigation potential” 

in 2030 is 1,382 x106 Mg C in 2030, or about 

46 x 106 Mg C/yr for the period 2000-2030. Th is 

assumes that forest restoration, conservation, and 

improved management programs continue, and that 

deforestation rates reach 25 percent of year-2000 level 

by 2030. Th is study also estimates potential costs of 

mitigation eff orts.

96. Masera, O.R.; Garza-Caligaris, J.F.; Kanninen, M.; 

Karjalainen, T.; Liski, J.; Nabuurs, G.J.; Pussinen, 

A.; de Jong, B.H.J.; Mohren, G.M.J. 2003. 

Modeling carbon sequestration in aff orestation, 

agroforestry and forest management projects: the 

CO2FIX V.2 approach. Ecological Modelling. 164: 

177-199.

Th is study describes and presents testing and validation 

results for version 2 of the CO2FIX ecosystem-level 

model (CO2FIX V.2), which is designed to estimate 

C balances of alternative management strategies for 

uneven- or even-aged forests in both temperate and 

tropical systems. Th e model estimates C stored in 

aboveground biomass, soils, and wood products. 

Testing and validation were conducted on an evenaged Picea abies monoculture in central Europe, an 

even-aged, mixed Fagus sylvatica and Pseudotsuga 

menziesii stand in Atlantic Europe, a mixed Pinus–

Quercus stand in central/southern Mexico, and a mixed 

cohort agroforestry plantation in Costa Rica. Th e 

ability to model multiple cohort stands and simulate 

both age- and biomass-based tree growth demonstrates 

the model’s fl exibility.

97. McCarney, G.R.; Armstrong, G.W.; Adamowicz, 

W.L. 2008. Joint production of timber, carbon,

and wildlife habitat in the Canadian boreal 

plains. Canadian Journal of Forest Research. 38: 

1478-1492.

An expansion of a previous eff ort is presented to model 

the tradeoff s associated with the joint production of 
29

timber supply and wildlife habitat. Th e previously 

developed standard linear programming model was 

expanded to include the Carbon Budget Model of the 

Canadian Forest Sector (CBM-CFS3). Th e result is a 

tool that allows for “extensive marginal cost analysis” 

of tradeoff s among production of C, timber, and 

wildlife habitat, and takes into account C in separate 

biomass and dead organic matter pools for individual 

forest-cover types.

98. McKenney, D.W.; Yemshanov, D.; Fox, G.; 

Ramlal, E. 2004. Cost estimates for carbon 

sequestration from fast growing poplar 

plantations in Canada. Forest Policy and 

Economics. 6: 345-358.

Th is study presents a model that estimates storage 

potential and break even points for aff orestation 

of marginal agricultural land in Canada based on 

prices for C sequestration and wood products and 

on agricultural-related opportunity costs. A variety 

of scenarios demonstrate that aff orestation is more 

economically attractive in western Canada, partially 

due to higher growth and productivity rates in this 

region.

99. Neilson, E.T.; MacLean, D.A.; Meng, F.R.; 

Arp, P.A. 2007. Spatial distribution of carbon 

in natural and managed stands in an industrial 

forest in New Brunswick, Canada. Forest Ecology 

and Management. 253: 148-160.

Th e Carbon Budget Model of the Canadian 

Forest Sector (CBM-CFS3) is used to simulate C 

accumulation in living biomass and dead organic 

material based on timber volume in major stand types 

in managed vs. unmanaged, as well as hardwood 

vs. softwood stands in northern New Brunswick, 

Canada. Species include Picea mariana, Picea rubens, 

Picea glauca, Abies balsamea, Pinus banksiana, Fagus 

grandifolia, Betula alleghaniensis, Acer saccharum, 

Populus tremuloides and Betula papyrifera. Th is paper 

contains extensive data on estimated mean total C 

density (Mg/ha) based on stand type, ecoregion, and 

age class for hundreds of stands. Total ecosystem C, 

excluding soil, is estimated to be 35 Mt over 428,000 

ha, or approximately 82 Mg C/ha. General fi ndings 

include: hardwoods (over 40 years) contain more C 

than similarly aged softwoods, by virtue of their greater 

wood density; total C in all biomass pools increased 

as a function of age, but total C accumulation varied 

according to species type; at 100 years, planted 

softwoods contained 94-135 Mg C/ha vs. 92-117 Mg 

C/ha for natural softwoods and 127-138 Mg C/ha for 

naturally regenerated hardwoods; C in most stands 

began to decline after 100 years, but much of this 

decline in older stands was related to an assumed stand 

break-up. Although simulation of C stocks was specifi c 

to a particular landscape, this modeling method 

(using merchantable volume to estimate C stock) can 

be applied more broadly allowing forest managers to 

optimize for timber values/harvesting relative to C 

sequestration.

100. Newell, R.G.; Stavins, R.N. 2000. Climate 

change and forest sinks: factors aff ecting the costs 

of carbon sequestration. Journal of Environmental 

Economics and Management. 40: 211-235.

Th is study examines the cost sensitivity of terrestrial 

C sequestration to relative prices, opportunity costs, 

and discount rates. Results show that the marginal 

cost of avoided deforestation (especially in the tropics) 

is signifi cantly lower than the cost of aff orestation/

reforestation and that discount rates are important 

in overall sequestration quantities. As discount rates 

increase, the marginal cost of sequestration increases 

monotonically and forestation also increases, but the 

present value of C sequestration per acre decreases.

101. Niu, X.; Duiker, S.W. 2006. Carbon 

sequestration potential by aff orestation of 

marginal agricultural land in the Midwestern 

U.S. Forest Ecology and Management. 223: 415-

427.

Th is study estimates sequestration potential through 

aff orestation of Midwest marginal agricultural land 

(Magland) based on forest type (coniferous vs. 

deciduous) and a simplifi ed management tradeoff 

(short-rotation vs. permanent forest). Th e study 

projects changes in aboveground biomass, root, 

forest fl oor, and soil organic C pools, and modifi ed 

sequestration rates based on management impacts 
30

and regional climate conditions such as temperature, 

precipitation, and site quality. Sequestration potential 

through Magland aff orestation ranges from 508-540 

Tg C over 20 years, and from 1018-1080 Tg C over 

50 years. Th ese ranges correspond to an estimated total 

of about 6.5 Mha Magland available in the United 

States. Coniferous forests show slightly greater overall 

potential in all scenarios, due to greater forest fl oor 

C accumulation (needles take longer to decompose 

than leaves). Overall mitigative potential would be 

approximately equal for both scenarios if “best C 

management” were used in the short rotation scenario, 

that is, harvested wood should be used for fossil off sets 

or as long-lived wood products. Otherwise, permanent 

forest would be slightly better for overall sequestration. 

Should aff orestation occur in these regions, slightly 

better C outcomes might result from regionally specifi c 

decisions with regard to species selected for planting. 

Magland aff orestation could off set roughly 7-8 percent 

of the region’s current GHG emissions over 20 years, 

or 6 percent over 50 years.

102. Palosuo, T.; Peltoniemi, M.; Mikhailov, A.; 

Komarov, A.; Faubert, P.; Th ürig, E.; Lindner, M. 

2008. Projecting eff ects of intensifi ed biomass 

extraction with alternative modeling approaches. 

Forest Ecology and Management. 255: 1423-1433.

Th e study uses two models (MOTTI-YASSO and 

EFIMOD-ROMUL) to simulate the eff ects of biomass 

extraction on forest C balances in six stands (three 

dominated by Pinus sylvestris and three by Picea abies) 

in Finnish forests. Th ere were substantial diff erences 

in predictions based upon the model used. Each 

had diff erent assumptions regarding the timing and 

intensity of thinnings and the rotation length/timing 

of fi nal harvest. Model outputs for soil C stocks and 

forest C stocks were similar for four of six stands, but 

harvest residue decomposition diff ered signifi cantly 

between models. Model comparison does not allow 

validation but it does allow an exploration of the 

behavior and theoretical limitations of the models and 

of the uncertainties associated with model structure.

103. Perez-Garcia, J.; Lippke, B.; Comnick, J.; 

Manriquez, C. 2005. An assessment of carbon 

pools, storage, and wood products market 

substitution using life-cycle analysis results. 

Wood and Fiber Science. 37: 140-148.

Results of a life cycle assessment of ho using production 

are used to create an accounting tool that “track[s] 

carbon from sequestration to substitution in forest 

product end-use markets.” Because substitution of wood 

for more fossil intensive products can have a signifi cant 

impact on reducing construction and housing-related 

emissions, management of forests for these products 

could contribute signifi cantly to mitigation.

104. Pohjola, J.; Valsta, L. 2007. Carbon credits and 

management of Scots pine and Norway spruce 

stands in Finland. Forest Policy and Economics. 9: 

789-798.

Th is study models a fi nancially optimum standlevel forest management scenario for Pinus sylvestris/

Picea abies stands in Finland, incorporating both 

timber and C values. Valuing C sequestration leads to 

higher stocking densities and longer rotation lengths. 

Th is fi nding is similar to those of other studies that 

compare traditional timber management scenarios with 

C-inclusive optimization models.

105. Pussinen, A.; Karjalainen, T.; Mäkipää, R.; 

Valsta, L.; Kellomäki, S. 2002. Forest carbon 

sequestration and harvests in Scots pine stand 

under diff erent climate and nitrogen deposition 

scenarios. Forest Ecology and Management. 158: 

103-115.

A forest succession applied gap-type model is used 

to simulate the eff ects of increased temperature, N 

deposition, and increased rotation length on timber 

and C sequestration in a simulated Pinus sylvestris

stand in southern Finland. Mean ecosystem C stock 

increased from 60 to 85 Mg C/ha when the rotation 

length was increased from 40 to 110 years. Th e rate 

of increase of C stock declined as rotation length 

increased (lengthening rotation from 40 to 45 years 

increased C stock by 3 Mg C/ha, while lengthening 

rotation from 90 to 95 years increased stock by 1 Mg 

C/ha) and changed climate conditions led to lower 

forest C stock due to decreased SOM. Other models 

suggest CO2 fertilization leads to greater ecosystem C 

accumulation.
31

106. Rokityanskiy, D.; Benítez, P.C.; Kraxner, F.; 

McCallum, I.; Obersteiner, M.; Rametsteiner, 

E.; Yamagata, Y. 2007. Geographically explicit 

global modeling of land-use change, carbon 

sequestration, and biomass supply. Technological 

Forecasting and Social Change. 74: 1057-1082.

Th is study reports on the results of a spatially explicit 

model, “Dynamic Integrated Model of Forestry and 

Alternative Land Use”, or DIMA, to quantify the 

economic potential for mitigation at both a domestic 

and international level for decisions on land-use 

management and biomass. DIMA integrates scenario 

planning by accounting for dynamic C and energy 

prices, global biomass supply curves, and global 

projections of land-use change, C, and bioenergy 

supply as it relates to socio-economic decision making 

and opportunity costs for alternative land uses. Th e 

model results are compared to those of similar studies. 

Th e costs of mitigation using DIMA are similar to 

other estimates at lower sequestration levels, but 

DIMA predicts lower-than-average costs at higher 

quantities (about $150/ton at the 6-7 Tg level, whereas 

Stavins (1999) predicts exponentially higher costs 

beyond 4-5 Tg). Results of this study found that C 

price incentives can generate signifi cant biodiversity 

and rural development cobenefi ts, and that C prices 

factor considerably into avoided deforestation eff orts; 

5-75 percent of global deforestation can be halted 

depending on C price. Th is study notes that the 

model should be expanded to account for risks and 

uncertainty (e.g., from fi re and pests).

107. Schmid, S.; Th ürig, E.; Kaufmann, E.; 

Lischke, H.; Bugmann, H. 2006. Eff ect of forest 

management on future carbon pools and 

fl uxes: A model comparison. Forest Ecology and 

Management. 237: 65-82.

Th e study compares empirical, semi-empirical, 

and process-based models to determine eff ects of 

four management scenarios (current harvest level, 

intensifi ed harvest, reduced harvest, and maximum 

sustainable growth harvest) on C pools and fl uxes in 

four even-aged forest types in Switzerland (coniferous, 

plateau; mixed deciduous, plateau; beech and fi r, preAlps; coniferous, Alps). Th e models predicted similar 

fl uxes, but there were some substantial diff erences 

in modeled C pools due in part to diff erent model 

assumptions and highly heterogeneous climate 

conditions. Results also show substantial potential for 

increased C storage, which emphasizes the need for 

full product accounting for accurate C accounting. 

Minimally harvested forests maximized C on-site 

but storage capacity was limited and peaked at 100 

years, after which the forests can become a C source. 

Th e “maximum sustainable growth/max carbon fl ow 

harvest” results in less on-site storage but has can 

continue mitigating beyond 100 years. Th e study 

concludes that no single strategy is “best” due to sitespecifi c conditions.

108. Scott, N.A.; Rodrigues, C.A.; Hughes, H.; Lee, 

J.T.; Davidson, E.A.; Dail, D.B.; Malerba, P.; 

Hollinger, D.Y. 2004. Changes in carbon storage 

and net carbon exchange one year after an initial 

shelterwood harvest at Howland Forest, ME.

Environmental Management. 33: S9-S22.

Th e study examines the changes in C storage and fl ux 

1 year after a shelterwood harvest on a red spruce 

and eastern hemlock-dominated forest in Maine. 

Preharvest C levels were 85 Mg C/ha; harvesting 

removed an estimated 14.9 Mg C/ha in timber, 

leaving 5.3 and 5.2 Mg C/ha in aboveground slash 

and root/stump detritus, respectively. About half of 

the harvested wood was used for long-lived products 

(estimated half-life of 45 years) and the other half for 

short-lived paper products (estimated half-life of 3.5 

years). Modeling estimated that the site is a net source 

of C until at least 5 years after harvest, indicating that 

shelterwood cuts remain a source for at least several 

years. Additional study and measurement of all C 

pools over longer time scales is required to determine 

when the site reverses to sink status and whether this 

type of management is optimal for this site.

109. Seely, B.; Nelson, J.; Wells, R.; Peter, B.; 

Meitner, M.; Anderson, A.; Harshaw, H.; Sheppard, 

S.; Bunnell, F.L.; Kimmins, H.; Harrison, D. 

2004. Th e application of a hierarchical, decisionsupport system to evaluate multi-objective 

forest management strategies: a case study in 
32

northeastern British Columbia, Canada. Forest 

Ecology and Management. 199: 283-305.

Th is study develops a framework for linking multiple 

forest management models within a hierarchy that 

facilitates data transfer, and uses it for multiple-use 

decision-making for a forest in British Columbia, 

Canada. Th e integrated decision-support system (DSS) 

allows tracking of soil, biomass, and dead organic 

pools under a natural disturbance baseline (control) 

and two alternative management approaches with 

higher harvesting levels. Carbon storage is relatively 

lower in higher harvesting scenarios, but storage 

declines in the baseline scenario, likely from C releases 

from fi re. Natural and climate-related disturbances 

should be integrated into DSS systems, which serve 

“at least four broad purposes: aiding research, guiding 

management, conveying knowledge, and facilitating a 

public evaluation of management tradeoff s.”

110. Seely, B.; Welham, C.; Kimmins, H. 2002. 

Carbon sequestration in a boreal forest 

ecosystem: results from the ecosystem simulation 

model, FORECAST. Forest Ecology and 

Management. 169: 123-135.

A silvicultural/forest management decision model, 

FORECAST, calibrated for a mixed wood forest 

(Picea glauca, Populus tremuloides, and Pinus contorta) 

is applied in British Columbia, Canada, to determine 

the C consequences of 11 management scenarios 

combining diff erent tree species, harvesting methods 

and rotation lengths, and fertilization over a 300-year 

period. Soil C stocks remain relatively stable regardless 

of treatment, aboveground biomass generally increases 

with rotation length, and wood product pool C 

“tended to decline at longer rotations.” Only the 150-

year pine and 200-year spruce rotations generate C 

storage that exceeded the natural disturbance scenario, 

though it is unclear whether the small additional 

gains were statistically signifi cant. Th e study notes 

declines in productivity over multiple rotations, but 

N fertilization counteracts this by increasing biomass 

accumulation by as much as 32 percent. Diff erent 

species are best suited to diff erent rotation lengths: 

aspen accumulates biomass quickly (1.6-1.8 Mg C/

ha/yr) and is ideal for short-rotation (<75 years); pine 

(1.3-1.7 Mg C/ha/yr) for mid-rotation (75-150 years); 

and spruce (1.0-1.4 Mg C/ha/yr) for longer rotations 

(150+ years). Th e results highlight the tradeoff 

between on-site ecosystem C storage and timber 

harvest, and as well as the importance of full product 

accounting for determining optimal rotation strategies. 

Th is study suggests managing stands for a variety of 

species and rotation lengths to provide for both timber 

and C sequestration values, or, alternatively, across a 

landscape to meet multiple criteria. Model outputs can 

help guide managers toward tree species and rotation 

lengths that are variably favorable for sequestration or 

timber.

111. Seidl, R.; Rammer, W.; Jäger, D.; Currie, W.S.; 

Lexer, M.J. 2007. Assessing trade-off s between 

carbon sequestration and timber production 

within a framework of multi-purpose forestry 

in Austria. Forest Ecology and Management. 248: 

64-79.

Th is study uses a hybrid model to determine the eff ects 

of three management strategies (age class forestry, 

90-year rotations; continuous cover forestry through 

structural thinnings, 90-year rotations; and moderate 

conversion to mixed broadleaf, 80-year rotations) on 

a Picea abies forest in Austria, taking into account 

wood product and bioenergy off set pools. Although 

the results demonstrate highest overall storage in the 

unmanaged (control) scenario (134.9 Mg/ha vs. 100.2 

Mg/ha for control and continuous cover management, 

respectively, over 100 years), results do not incorporate 

the uncertainty and risks associated with alternative 

strategies. For example, reductions generated through 

fossil fuel substitution are permanent and real, whereas 

standing C on the landscape (unmanaged/control) 

is always subject to unforeseen disturbance-related 

reversals in C storage. Th is study also notes that forestbased C sequestration is cost-effi cient, but has limited 

biological and societal potential.

112. Seidl, R.; Rammer, W.; Jäger, D.; Lexer, M.J. 

2008. Impact of bark beetle (Ips typographus L.) 

disturbance on timber production and carbon 

sequestration in diff erent management strategies 

under climate change. Forest Ecology and 

Management. 256: 209-220.
33

Th is study models the combined impact of bark beetle 

disturbance and climate changes on timber production 

and C sequestration in secondary Picea abies stands 

in Austria that already were outside their natural 

range and thus even more vulnerable to threats. Four 

management alternatives/strategies (age class forestry, 

continuous cover forestry, conversion to mixed species 

stands, and no management) for Picea abies stands 

were paired with one baseline and two climate change 

scenarios (ECHAM4-OPYC3 and HadCM2), and two 

disturbance scenarios (no disturbance and bark beetle 

damage). Beetle damage is far more severe (increases of 

219 percent in timber losses) for the changed climate 

vs. baseline scenario. Paired climate change and 

disturbance scenarios resulted in signifi cantly reduced 

C storage in managed stands (up to 41 Mg/ha) 

whereas some unmanaged stand simulations showed 

slight increases due to stand density eff ects. When 

climate change and beetle disturbances are included, 

continuous cover forestry achieves higher ecosystem C 

gains than other active management strategies. When 

bark beetle disturbance is eliminated, all managed 

stand variants fared better in terms of timber stocks 

than unmanaged stands in climate change scenarios.

113. Sivrikaya, F.; Keleş, S.; Çakir, G. 2007. Spatial 

distribution and temporal change of carbon 

storage in timber biomass of two diff erent forest 

management units. Environmental Monitoring 

and Assessment. 132: 429-438.

Spatially explicit estimates and mapping of C storage can 

help link aboveground and belowground processes and 

help land managers make more informed decisions; 

this paper proposes a methodology to create spatially 

explicit carbon estimates. Th e study maps C storage 

for two diff erent forest ecosystems, and evaluates the 

results in the context of changes in land use, forest 

cover, and species-specifi c growing stocks.

114. Spring, D.; Kennedy, J.; Mac Nally, R. 2005. 

Optimal management of a fl ammable forest 

providing timber and carbon sequestration 

benefi ts: an Australian case study. Th e Australian 

Journal of Agricultural and Resource Economics. 

49: 303-320.

Extending rotation length often is considered a 

pragmatic option for enhancing sequestration. Th is 

study uses stochastic dynamic programming to explore 

and introduce wildfi re risk (and associated unintended 

C releases) into risk analysis, thinning, and harvesting 

decisions over a range of potential C prices. Th e eff ect 

of treating saw logs as permanent post-harvest sinks is 

also analyzed. As C exceeds AU $60/ton, the optimal 

planned harvest age decreases.

115. Sohngen, B.; Brown, S. 2006. Th e infl uence 

of conversion of forest types on carbon 

sequestration and other ecosystem services in 

the South Central United States. Ecological 

Economics. 57: 698-708.

More than 12 million acres of agricultural land in 

the south-central United States have been shifted 

into forest plantations over the past 40 years. Th is 

study examines potential shifts in land-use as a result 

of increased demand/prices for timber products, 

land rental rates, and agricultural commodity prices. 

Modeled results show that future pine plantations 

likely will be established by converting natural 

forests, resulting in reduced capacity for C storage by 

as much 0.9 Tg C/yr in aboveground biomass and 

harvested wood products in Arkansas, Louisiana, and 

Mississippi. Natural forests and associated greater C 

storage might be maintained through subsidies.

116. Stinson, G.; Freedman, B. 2001. Potential 

for carbon sequestration in Canadian forests 

and agroecosystems. Mitigation and Adaptation 

Strategies for Global Change. 6: 1-23.

Canadian forests currently are a net source of 

emissions. A large though ultimately limited land area 

and the risk of catastrophic C losses from disturbances 

limit the amount of C that can be stored in Canada’s 

forest ecosystems, but long-term opportunities for 

mitigation exist through fossil fuel displacement. 

Th is study examines the potential for optimizing 

mitigation in Canadian agroecosystems and in four 

forest ecosystems: boreal aspen, boreal white spruce, 

coastal Douglas fi r, and Acadian mixed tolerant 

hardwoods through four management strategies: 

maximum sustained yield (MSY) management for 
34

pulp logs, MSY management for saw logs, MSY 

management for biofuel, and creation of protected 

forest C reserves. Th e C storage potential of each of 

the four strategies for each ecosystem is discussed. In 

all ecosystem types, total mitigation is greatest after 

200 years with the protected C reserve strategy (131 

Mg C/ha for boreal aspen, 161 Mg C/ha for boreal 

white spruce, 473 Mg C/ha for coastal Douglas fi r, 

and 94.7 Mg C/ha for Acadian mixed hardwoods). 

However, mitigation strategies that entail displacement 

of more energy-intensive products are irreversible 

and thus more certain (not subject to catastrophic 

loses of C from the landscape). Th is untitled “simple 

budget model” (which takes into account aboveground 

and belowground C, harvested wood products, 

and displaced emissions) assumes suppression of 

“natural” disturbances, which typically occur on 30-

200 year intervals. In reality, however, suppressing 

natural disturbances could be costly, risky, and/or 

counterproductive (e.g., fi re suppression could lead 

to more catastrophic fi res). As a result, it is worth 

considering the benefi ts of other management options, 

particularly MSY management for saw logs, which 

resulted in C storage that was nearly as substantial.

117. Th ürig, E.; Palosuo, T.; Bucher, J.; Kaufmann, 

E. 2005. Th e impact of windthrow on carbon 

sequestration in Switzerland: a model-based 

assessment. Forest Ecology and Management. 

210: 337-350.

Although most forests act as important C sinks, 

disturbances such as windstorms can result in 

premature and potentially large C releases, especially in 

older forests. Th is study examines the eff ects of wind 

disturbance on Swiss forests, typically dominated by 

Picea abies, Fagus, and Abies alba, using two validated 

models: a forest management and a C model. Th is 

study combines two management scenarios (business 

as usual [BAU] and reducing large logs [RLL]), two 

storm scenarios, and two post-storm prescriptions 

for a total of 10 management-disturbance scenarios 

and fi nds that: BAU management scenarios result 

in substantial accumulations of C on the landscape, 

whereas RLL management led to a negative total C 

budget after 30 years. Forest soil C increases are slight 

and slow; and that post-disturbance clearing only 

slightly reduces the C budget. A 30 percent increase 

in the frequency of storms had “only a small impact 

on carbon sequestration in forests” in part because 

the input of windthrown timber to soil C can more 

than off set the C released from uprooting. A “best” 

mitigation strategy cannot be recommended without 

accounting for long-term soil C eff ects and the use of 

salvaged wood.

118. Trofymow, J.A.; Stinson, G.; Kurz, W.A. 

2008. Derivation of a spatially explicit 86-year 

retrospective carbon budget for a landscape 

undergoing conversion from old-growth to 

managed forests on Vancouver Island, BC. Forest 

Ecology and Management. 256: 1677-1691.

Th e study employs the CBM-CFS3 model along 

with historical disturbance maps, ortho-photographs, 

and a 1919 timber cruise map to reconstruct the 

C budget for the Pseudotsuga menziesii-dominated 

Oyster River area in eastern Vancouver Island in 

British Columbia, Canada, to determine C-related 

eff ects of conversion from an old-growth to managed 

forest. It was diffi cult to construct a C budget due to 

the diffi culty of reconstructing the initial condition 

of the forest. Th e study estimates total aboveground, 

belowground, soil and total C storage of 271, 55, 237, 

and 749 Mg C/ha in 1920 (prior to logging) which is 

similar to modern-day total ecosystem storage in H.J. 

Andrews Experimental Forest in Oregon (625 Mg C/

ha) and Olympic National Park in Washington (819 

Mg C/ha). Not surprisingly, C stocks declined from 

1930-1945 when the forest was subjected to fi res 

and logging. Th e forest remained a C source until 

1954, losing an average of 365-411 Mg/ha for a total 

of 0.91-1.02 Mt C. Harvesting of second growth 

in the 1980s again reduced the C budget but stocks 

recovered to 504-533 Mg C/ha by 1998. “Despite 

their high productivity, the area’s forests are not 

likely to obtain C densities of the landscape prior to 

industrial logging because the stands will not reach 

pre-logging ages.”

119. Van Wijk, M.T.; Dekker, S.C.; Bouten, W.; 

Kohsiek, W.; Mohren, G.M.J. 2001. Simulation of 

carbon and water budgets of a Douglas-fi r forest. 

Forest Ecology and Management. 145: 229-241.
35

A forest growth-hydrology model (FORGRO-SWIF) 

that measures water-use effi ciency was validated 

and used to model forest water use for thinned and 

unthinned stands in a Pseduotsuga menziesii forest in 

the Netherlands. Th is study fi nds the C changes in the 

unthinned stand are double the C sink of the thinned 

stand (10-12.1 Mg C/ha/yr vs. 4.3 -6.2 Mg C/ha/yr). 

Water-use effi ciency is higher in the unthinned scenario.

120. Vetter, M.; Wirth, C.; Bottcher, H.; Churkina, 

G.; Schulze, E.D.; Wutzler, T.; Weber, G. 2005. 

Partitioning direct and indirect human-induced 

eff ects on carbon sequestration of managed 

coniferous forests forests using model simulations 

and forest inventories. Global Change Biology. 11: 

810-827.

Th is study attempted to isolate the eff ects of indirect 

human-driven changes (N deposition and CO2 

fertilization) and forest age class distribution and legacy 

on the total C sink of a Picea abies/Pinus sylvestrisdominated coniferous forest in Germany. Th e study 

uses the BIOME-BGC model to simulate C, N, water 

fl uxes, forest age dynamics, and site history and fi nds 

that: mature forests (>70 years) are signifi cant C sinks 

of up to 5 Mg C/ha/yr; mature and young stands both 

show signifi cant responses to N deposition, whereas 

middle-aged stands were already growing at their 

maximum rate and did not exhibit much additional 

response; and CO2 fertilization acts to improve 

water-use effi ciency and drives greater biomass gains, 

especially at low and medium elevation sites. Indirect 

human infl uences result in a 33 percent increase in 

forest productivity and the age-class distribution 

resulting from previous management decisions accounts 

for roughly 8-17 percent of the current sink. Findings 

diff er signifi cantly from Caspersen et al. (2000), who 

concluded that indirect human impacts play a much 

smaller role (2 percent) in increasing forest productivity 

based on repeated forest inventory data.

III. Experiments with Some Pools 

Modeled

121. Bauer, G.A.; Bazzaz, F.A.; Minocha, R.; 

Long, S.; Magill, A.; Aber, J.; Berntson, G.M. 

2004. Eff ects of chronic N additions on tissue 

chemistry, photosynthetic capacity, and carbon 

sequestration potential of a red pine (Pinus 

resinosa Ait.) stand in the NE United States. 

Forest Ecology and Management. 196: 173-186.

Results from a 15-year study of N amendments on 

Pinus resinosa on the Harvard forest in Massachusetts 

demonstrate that while N amendments have 

signifi cantly increased foliar N content, the 

treatment reduced foliar longevity and decoupled the 

photosynthesis-N relationship resulting in decreased 

photosynthetic capacity (of fertilized trees) by 50 

percent. An increase in N foliar content does not 

correspond to a greater capacity for C assimilation 

primarily because increased N reduces foliar longevity. 

Authors used the PnET-Day model to determine 

eff ects on long-term C balance; simulation results are 

vague and general, suggesting a reduced net C gain.

122. Banfi eld, G.E.; Bhatti, J.S.; Jiang, H.; Apps, 

M.J. 2002. Variability in regional boreal forest 

ecosystems: results from West-Central Alberta.

Forest Ecology and Management. 169: 15-27.

Th is study uses a combination of forest inventory data, 

model simulations, fi eld-observed plot data, and soilpolygon information to model regional estimates and 

spatial variation in aboveground biomass, forest fl oor, 

and soil C stocks for a transitional boreal region in 

Alberta, Canada that is dominated by Pinus contorta

with Populus tremuloides and Picea glauca. Th e current 

regional average aboveground biomass C was estimated 

at 43 Mg C/ha. Th e model revealed the major role 

of natural and human disturbance from 1920-1995. 

Improved forest C budget models can better help track 

management and climate-induced changes.

123. Hoover, C.; Stout, S. 2007. Th e carbon 

consequences of thinning techniques: Stand 

structure makes a diff erence. Journal of Forestry. 

105: 266-270.

Th inning often is used to increase the quality and size 

of merchantable timber. Th is study examines stands 

that have been thinned twice over 25 years using three 

techniques (thinned from below, 60-70 percent target 

density; thinned from middle, 60-70 percent target 

density; and thinned from above). Th e study fi nds a 

net C benefi t of 0.15 Mg C/ha/yr (in terms of live 
36

aboveground biomass, slash, deadwood, and products) 

in stands that were thinned from below compared 

to the control. Stands thinned from middle or above 

exhibit net C losses of 0.99 Mg C/ha/yr and 1.41 Mg 

C/ha/yr, respectively, as compared to the control.

124. Kashian, D.M.; Romme, W.H.; Tinker, 

D.B.;Turner, M.G.; Ryan, M.G. 2006. Carbon 

storage on landscapes with stand-replacing fi res.

BioScience. 56: 598-606.

Th e eff ects of stand-replacing fi res on short and longerterm landscape C storage are examined for coniferous 

forests in light of fi re disturbance becoming more 

frequent and more likely due to historical and climaterelated factors. Near-term eff ects of wildfi re on forest 

ecosystem C balance were considered, including the 

balance between C loss in deadwood decomposition 

and C gain in new vegetation. Longer-term C storage 

trajectories for a landscape are related to regeneration 

profi les. Changing fi re frequency aff ects stand age 

structure and increasing frequency changes the age 

distribution of forest stands, but equilibrium values 

of landscape C storage are resistant to change unless 

fi re intervals become substantially shorter. Th e 1988 

Yellowstone wildfi res were used to explore near and 

longer-term C dynamics. Results showed direct fi rerelated C losses of 13.6 Mg C/ha; 15.3 Mg C/ha 

will be lost through decomposition over 250 years 

following the 1988 fi re; that the landscape will return 

to prefi re NEP levels by 2028; and that total C lost 

will be recovered in 230 years.

125. Martin, J.M.; S.T.G., Plaut, J.; Holmes, B. 2005. 

Carbon pools in a boreal mixedwood logging 

chronosequence. Global Change Biology. 11: 

1883-1894.

Th is study examines aboveground carbon and 

detritus in a mixedwood boreal forest (dominated 

by Betula papyrifera, Larix laricina, Picea mariana, 

Picea glauca, Pinus banksiana, Populus balsamifera, 

Populus tremuloides) in Manitoba, Canada. Th e 

eff ects of harvesting on C in these forests had not 

been studied previously, although previous studies 

used the BOREAS model, which was designed for 

relatively pure boreal forest types. Four even-aged 

mixed-wood stands that were 11, 18, 30, and 65 

years old are considered, the fi rst three of which 

originated following a clearcut and the fourth 

originated following wildfi re. C density in these 

mixedwood stands is 47 percent higher than a pure 

wet Picea mariana stand, and 44 percent higher than 

a pure Pinus banksiana stand (all boreal). Possible 

explanations include: litterfall, forest fl oor, and tree N 

content are greater for deciduous vs. coniferous forests; 

natural succession in a boreal forest ecosystem includes 

a transition from deciduous to coniferous trees, 

and canopy production effi ciency and transpiration 

declines during this succession; and/or that mixedwood stands have multilayered canopies that can 

support greater foliage mass.

126. McNulty, S.G. 2002. Hurricane impacts on 

U.S. forest carbon sequestration. Environmental 

Pollution. 116: S17-S24.

Hurricanes can be a major disturbance factor on 

forests. Ultimate impacts on the United States C 

budget depend on how much dead biomass results 

from a typical storm, the fate of the dead biomass 

(whether it is allowed to decay, used for bioenergy or 

wood products) and whether hurricanes confer longterm benefi ts on forests, e.g., increased nutrient inputs 

that may result in increased productivity. Th e increase 

in the detritus pool from four major hurricanes 

during the 20th century ranged from 2.9 to 20 Tg C. 

Results indicate that a single hurricane can “convert 

the equivalent of 10 percent of the total annual 

U.S. forest carbon sequestration from living to dead 

wood.” Although some downed wood is harvested 

after storms, much remains inaccessible. Hurricane 

damage can increase wildfi re risk and longer-term 

C losses associated with leaf area recovery time and 

reduced stem stocking probably are not off set by shortterm increases in forest. In hurricane-prone areas of 

the southeast, managers can mitigate against future 

storm damage by planting more longleaf pine and 

hardwoods, both of which are more storm resistant 

than other pines.

127. Radtke, P.J.; Amateis, R.L.; Prisley, S.P.; 

Copenheaver, C.A.; Chojnacky, D.C.; Pittman, 

J.R.; Burkhart, H.E. 2009. Modeling production 
37

and decay of coarse woody debris in loblolly pine 

plantations. Forest Ecology and Management. 257: 

790-799.

Th e volume, density, and mass of CWD at 36 Pinus 

taeda plots throughout the southeastern United States 

were sampled. Th e plots were installed in 21-yearold stands that remained from 186 stands established 

in 1980-82 to study thinning eff ects. Data on tree 

mortality data from the original study were used to 

create decay classes for dead trees. Th ese were paired 

with on-site CWD data to create prediction equations 

for CWD production, decay, and yield. Th e equations 

can predict CWD yield at the tree to plantation level, 

and can estimate CWD C stocks for loblolly pine 

plantations throughout the southeast.

128. Ryan, M.G.; Binkley, D.; Fownes, J.H.; 

Giardina, C.P.; Senock, R.S. 2004. An

experimental test of the causes of forest growth 

decline with stand age. Ecological Monographs. 

74: 393-414.

Th is study seeks to determine the reason for a 

decline in aboveground wood production (and 

thus C production) after canopy closure in stands 

of Eucalyptus saligna in Hawaii. Aboveground and 

belowground C budgets were measured for 6 months 

to 6.5 years after planting. Two stocking density levels 

(10,000 and 1,111 trees/ha) and three fertilization 

treatments were included. Meteorological and 

ecophysiological data were used to estimate C storage 

and fl ux, foliage and wood respiration rates, foliar 

nutrients, photosynthetic capacity, and other variables 

to determine whether the decline in aboveground 

wood/C production decline was related to a: 1) 

partitioning shift from wood production to respiration; 

2) proportional reduction in GPP driven by nutrient 

limitation, reduced canopy leaf area/photosynthesis, 

increased abrasion between tree canopies, or other 

reasons; or 3) combination of these. Results support 

the combination hypothesis: GPP declined and the 

decline in wood/C production was proportionally 

greater than the decline in canopy photosynthesis.

129. Sampson, D.A.; Waring, R.H.; Maier, C.A.; 

Gough, C.M.; Ducey, M.J.; Johnsen, K.H. 2006. 

Fertilization eff ects on forest carbon storage and 

exchange, and net primary production: A new 

hybrid process model for stand management. 

Forest Ecology and Management. 221: 91-109.

A hybrid model was calibrated and validated to predict 

the eff ects of N fertilization on Pinus taeda stands 

growing in nutrient poor soils of North Carolina. N 

fertilization was used to increase leaf-area index and 

NEP by tenfold over the control treatment (0.73 

Mg C/ha/yr over a 12-year period). Seasonal uptake 

patterns suggested that “autumn and winter may 

be critical periods for C uptake in nutrient-limited 

[Pinus taeda] stands.” However, accumulation of 

aboveground biomass was 27 percent less than that 

found by Jokela and Martin (2000) in intensively 

managed loblolly pine stands in Florida. Th e C 

impacts of intensively managed stands depend heavily 

on management and its long-term eff ect on soil C 

and on whether harvested wood is made into shortor long-lived products; neither of these factors was 

included in this study.

130. Taylor, A.R.; Wang, J.R.; Kurz, W.A. 2008. 

Eff ects of harvesting intensity on carbon stocks 

in eastern Canadian red spruce (Picea rubens) 

forests: An exploratory analysis using the CBMCFS3 simulation model. Forest Ecology and 

Management. 255: 3632-3641.

Th is study compares results from a C budget model to 

actual C stocks and changes in Picea rubens dominated 

stands in Nova Scotia, Canada. Simulations of partial 

and clearcut harvests over 240 years were run to 

determine eff ects on C stocks. Although the partial 

harvest resulted in more C retained on the landscape 

than the clearcut (327.3 Mg C/ha vs. 305.8 Mg C/ha), 

the latter generated more merchantable timber (132.4 

Mg C/ha vs. 115.6 Mg C/ha). Th e partial-cut scenario 

resulted in more DOM C, likely the result of greater 

DOM inputs from selective harvesting methods. 

Additional research is needed to determine harvesting 

scenarios that would optimize timber output along 

with landscape C storage.

131. Th ornton, P.E.; Law, B.E.; Gholz, H.L.; Clark, 

K.L.; Falge, E.; Ellsworth, D.S.; Goldstein, A.H.; 
38

Monson, R.K.; Hollinger, D.; Falk, M.; Chen, J.; 

Sparks, J.P. 2002. Modeling and measuring the 

eff ects of disturbance history and climate on 

carbon and water budgets in evergreen needleleaf 

forests. Agricultural and Forest Meteorology. 113: 

185-222.

Th e eff ects of disturbance history and climate change 

on C and water fl uxes are reported for seven evergreen 

coniferous forests in California, North Carolina, 

Florida, Maine, Oregon, Colorado, and Washington 

using data and fi ndings from observations of canopy 

fl ux with comparisons to modeled eff ects. Th e unique 

disturbance history of each site (ranging from clearcut 

to selective harvest to low-level and stand-replacing 

fi res) is described. Results indicate that variations 

between sites are primarily related to disturbance 

history while other site-specifi c and changing N 

deposition and atmospheric CO2 concentrations play 

a secondary role. In all cases, predicted peak emissions 

related to disturbance occur within 2 years of 

disturbance, after which each site continues as a source 

of emissions for 4-16 years and then becomes a strong 

and gradually diminishing sink. Th e eff ects of CO2 

fertilization are limited by nutrient availability but 

eff ects are greatest following disturbance when nutrient 

availability is high and plant demand is relatively low.

IV. Literature Reviews and 

Meta-analyses

132. Birdsey, R.A.; Alig, R.; Adams, D. 2000. 

Mitigation activities in the forest sector to reduce 

emissions and enhance sinks of greenhouse 

gases. In: Joyce, L.A.; Birdsey, R.A., eds. Th e 

impact of climate change on America’s forests: a 

technical document supporting the 2000 USDA 

Forest Service RPA assessment. Gen Tech. Rep. 

RMRS-59. Ft. Collins, CO: U.S. Department 

of Agriculture, Forest Service, Rocky Mountain 

Research Station: 112-131.

Th is chapter provides a high-level analysis of 

opportunities to reduce sources (substitute wood 

products for more energy-intensive products; reduce 

the energy required to grow, harvest, and process 

timber; reduce the risk of catastrophic wildfi re), 

enhance sinks (aff orest land; avoid deforestation; 

improve management; reduce harvest/extend rotation; 

encourage agroforestry), and achieve concurrent source 

reductions/sink enhancements of GHGs (substitute 

bioenergy for fossil-based energy; increase C capture 

and retention in wood products; increase paper and 

wood reuse/recycling; enhance urban forestry/tree 

plantings). Th e estimated increase in U.S. forest C 

stocks from 1952-1992 was 281 Tg C/yr. Th e study 

estimates reduced future rates of storage (2000-

2040), on the order of 183 Tg C/yr. A summary 

table of enhanced C storage opportunities by practice 

is included. Additional opportunities that span the 

aforementioned options for reducing sources and/or 

simultaneously enhancing sinks are estimated at 210 

Tg C/yr.

133. Birdsey, R.; Pregitzer, K.; Lucier, A. 2006. Forest

carbon management in the United States: 1600-

2100. Journal of Environmental Quality. 35: 1461-

1469.

Th is study explains how past land-use practices in 

the United States have and will continue to aff ect 

the forest C budget. From 1600-1800, U.S. forests 

were in C balance with the atmosphere. Th ese forests 

have been a signifi cant sink in recent decades due to 

regrowth after the abandonment of agricultural land. 

Th e strength of the sink is declining but additional 

sequestration of 100-200 Tg C/yr may be possible 

through a combination of aff orestation, mineland 

reclamation, forest restoration and management, 

agroforestry, and the use of bioenergy and wood 

products. Th is will require improved monitoring and 

technology transfer to land managers.

134. Conant, R.T.; Paustian, K.; Del Grosso, S.J.; 

Parton, W.J. 2005. Nitrogen pools and fl uxes 

in grassland soils sequestering carbon. Nutrient 

Cycling in Agroecosystems. 71: 239-248.

Th e study demonstrates that anywhere from 10-

125 percent of C sequestration gains due to changes 

in management practices on grasslands were off set 

by increases in N2O fl ux. Conversely, altering 

management to minimize N2O fl ux may signifi cantly 

benefi t climate change mitigation eff orts. Some studies 

have shown that N sequestration and decreased N 
39

mineral availability lead to reduced N2O fl ux though 

this result was not corroborated by this study. Authors 

have demonstrated that N fertilization, relatively 

common on forests and grasslands, does not always 

increase C sequestration. Production, transport, 

and increased N2O fl ux that may be associated with 

fertilizer application can increase both CO2 and N2O 

emissions.

135. Gielen, B.; Ceulemans, R. 2001. Th e likely 

impact of rising atmospheric CO2 on natural 

and managed Populus: a literature review. 

Environmental Pollution. 115: 335-358.

Th is publication is a comprehensive review of literature 

pertaining to CO2 enrichment studies, including 

those related to controlled environment and open-top 

chambers, natural springs, and FACE experiments. 

Th ere is an obvious response by aboveground and 

belowground biomass of Populus spp. to CO2 

enrichment: responses for total or stem biomass ranged 

from 22-90 percent increased productivity (mean 

= 33 percent); leaf area increased by 7-103 percent; 

and root mass in both potted and fi eld experiments 

generally increased, though long-term relative growth 

response to CO2 fertilization remains unclear and 

is not necessarily greater than for slower growing 

species. Forest management (especially plantation 

management) can be modifi ed (rotation lengths 

shortened) to take advantage of greater productivity. 

Wood quality and vulnerability of certain poplar 

species to fungus, insects, and disease under changed 

climate conditions must be considered carefully.

136. Gower, S.T. 2003. Patterns and mechanisms 

of the forest carbon cycle. Annual Reviews Energy 

and Environment. 28: 169-204.

Th is publication provides an accessible general 

overview of the forest C cycle and the eff ects of 

climate-related disturbances and anthropogenic 

perturbations on C balances and exchanges of CO2

with the atmosphere. Th e forest C cycle is explained 

in relation to NPP, soil surface CO2 fl ux, and NEP. 

Disturbances related to climate change and their 

eff ects on forest ecosystems are summarized; a brief, 

high-level explanation of C sequestration in forest 

vegetation, soils, and forest products as well as off sets 

related to fossil fuel substitution is provided.

137. Han, F.; Plodinec, M.; Su, Y.; Monts, D.; Li, Z. 

2007. Terrestrial carbon pools in southeast and 

south-central United States. Climatic Change. 84: 

191-202.

Th is study estimates current annual rates of C 

sequestration in Alabama, Arkansas, Florida, Georgia, 

Louisiana, Mississippi, North Carolina, South 

Carolina, Tennessee, Texas, and Virginia, develops 

estimates for regional sinks, and determines how much 

of the region’s annual emissions might be off set by 

terrestrial sequestration. However, the study relied on 

1990 regional GHG emission estimates (more recent 

estimates were not available at the time), and on forest 

C pool estimates from 1996 and growth rates from 

1999. Th e study estimates that terrestrial sinks capture 

23 percent of total regional (southeast/south-central) 

GHG emissions and that total sink size in this region is 

130 Tg C/yr (more than 158,300 ha).

138. Heath, L.S.; Smith, J.E. 2000. An assessment of 

uncertainty in forest carbon budget projections. 

Environmental Science & Policy. 3: 73-82.

Th is study analyzes uncertainty forecasts for forest C 

inventory and fl ux. For inventory, uncertainty estimates 

were ±9 percent for 2000, increasing to 11 percent in 

2040. Th e greatest sources of uncertainty are soil and 

tree C for inventory estimates. Uncertainties related 

to fl ux estimates are “higher and more variable” than 

stock uncertainties, and greatest sources of uncertainty 

are related to volume growth and volume removals. 

Diff erent approaches are required to reduce uncertainty 

in fl ux estimates compared to those that are required to 

reduce uncertainty in stock (inventory) estimates.

139. Houghton, R.A. 2003. Why are estimates of 

the terrestrial carbon balance so diff erent? Global 

Change Biology. 9: 500-509.

Author reviews recent top-down and bottom-up 

estimates of the global C sink and sorts by source and 

decade. A northern mid-latitude sink of 2 Pg/yr is 

estimated primarily as a result of historical land-use 

change.
40

140. Houghton, R.A.; Hackler, J.L. 2000. Changes 

in terrestrial carbon storage in the United States. 

1: Th e roles of agriculture and forestry. Global 

Ecology & Biogeography. 9: 125-144.

Th is study examines ways in which historical land-use 

patterns have aff ected the U.S. C fl ux, provides estimates 

of fl ux magnitude and direction for past years based 

on these changes, and compares the U.S. C sink with 

atmospheric modeling of the North American C sink.

141. Houghton, R.A.; Hackler, J.L.; Lawrence, K.T. 

2000. Changes in terrestrial carbon storage in 

the United States. 2: Th e role of fi re and fi re 

management. Global Ecology & Biogeography. 9: 

145-170.

Th is is a second in a series of chapters that 

examines historical change in the U.S. C profi le. In 

examining the role of fi re on terrestrial C storage and 

accumulation, this study estimates the changes in land 

use (from forest to agricultural land) that reduced 

fi re incidence as well as the eff ects of fi re suppression. 

Th e study hypothesizes that this allowed for woody 

encroachment into some areas. Results show a total 

U.S. terrestrial sink of 390 Tg C/yr from 1950-1990, a 

signifi cant portion of which is related to fi re exclusion 

and fi re management. Th is paper contains much data 

related to historical burn intervals and current and 

historical terrestrial C accumulation in U.S. regions as 

a result of fi re management.

142. Hu, H.; Wang, G.G. 2008. Changes in forest 

biomass carbon storage in the South Carolina 

Piedmont between 1936 and 2005. Forest Ecology 

and Management. 255: 1400-1408.

Researchers examined changes in C stock in South 

Carolina’s Piedmont region. Conversion of 2.8 Mha of 

land from agricultural use to forest land between 1936 

and 2005 resulted in a net C gain of 1.19 Tg C/yr, or 

about 0.43 Mg C/ha/yr. Eighty percent of these gains 

were in aboveground biomass. Although soil C usually 

constitutes at least half of the total forest C pool, its 

soil content rarely is subject to rapid change.

143. Hyvonen, R.; Agren, G.I.; Linder, S.; Persson, T.; 

Cotrufo, M.F.; Ekblad, A.; Freeman, M.; Grelle, A.; 

Janssens, I.A.; Jarvis, P.G. 2007. Th e likely impact 

of elevated CO2, nitrogen deposition, increased 

temperature and management on carbon 

sequestration in temperate and boreal forest 

ecosystems: a literature review. New Phytologist. 

173: 463-480.

Th is study summarizes single-factor climate feedback 

fi ndings and explores the combined eff ects of 

increasing CO2, temperature, and N deposition, and 

intensifi ed management. Th e study states that: 1) 

hundreds of studies have shown that elevated CO2 

concentration stimulates tree growth but that longterm eff ects remain uncertain, including whether 

growth will be restricted by nutrient availability; 

2) single-factor studies have found that elevated 

temperature increases photosynthetic rates, CO2 and 

temperature frequently interact, and although leafscale photosynthetic rates may increase, increases in 

leaf-area may create more stand shade and counteract 

productivity increases; 3) decomposition of leaf litter 

increases with increasing temperature but the eff ect of 

temperature/response of leaf litter becomes less clear 

as the litter reaches later stages of decomposition; 4) 

soil respiration may briefl y increase, but a long-term 

eff ect is unlikely; 5) N deposition likely will increase C 

sequestration in nutrient limited environments; 6) the 

eff ects of N deposition on soil remain uncertain but a 

recent meta-analysis suggests that litter decomposition 

is stimulated at sites with low ambient N deposition 

(<5 kg/ha/yr) and high-quality litters, but that sites 

with moderate N deposition or low-quality litters 

experience reduced decomposition rates (5-10 kg/ha/

yr). Th e study also includes a table that summarizes 

the qualitative eff ects of various management practices 

and disturbance events on soil, biomass, and ecosystem 

C stock. Ecosystem C stocks generally increase with 

low-moderate site preparation, a change in species 

composition from broadleaves to conifers, fertilization, 

and increased rotation lengths. Ecosystem C stocks 

decrease with changes in species composition from 

conifers to broadleaves, and with some thinning 

methods. Ecosystem C may increase, decrease, or 

remain the same with: prescribed burning, peatland 

drainage, intensive site preparation, and harvesting 

method used.
41

144. Jones M.B.; Donnelly, A. 2004. Carbon 

sequestration in temperate grassland ecosystems 

and the infl uence of management, climate and 

elevated CO2. New Phytologist. 164: 423-439.

Th is is a thorough overview of the dynamics of C 

storage in temperate grassland ecosystems (ecosystems 

in which herbaceous species are the dominant 

vegetation) including a review of the literature on 

the eff ects of management and climate change on 

grassland C sequestration. Th e accumulation of C in 

grasslands is controlled largely by site productivity; 

estimates of when the soil C pool will reach a 

saturation point for temperate grasslands range from 

10-100 years. Management actions to increase C 

storage on grasslands include fertilization, irrigation, 

intersowing of grasses and legumes, and introducing 

productive forage grasses. It is diffi cult to determine 

the long-term eff ects of elevated concentrations of 

CO2 on grasslands: elevated CO2 concentrations 

often lead to increases in NPP of grasslands and 

much uncertainty remains with respect to respiration, 

turnover, C root allocation, and interaction with other 

nutrient demands. Sequestration rates for temperate 

pasture range from 0 to 8 Mg/ha/yr; improved 

management of U.S. pastureland may increase 

sequestration by 0.2-0.5 Mg C/ha/yr.

145. Litton, C.M.; Raich, J.W.; Ryan, M.G. 2007. 

Carbon allocation in forest ecosystems. Global 

Change Biology. 13: 2089-2109.

Th is meta-analysis reviews of 63 studies to determine 

the limits, patterns, and plasticity of biomass, fl ux, 

and partitioning—three critical components of C 

allocation. Nine broad generalizations are presented, 

including: biomass should not be used to infer fl ux 

or partitioning, increased GPP increases C fl ux of all 

ecosystem components, and C fl uxes vary more than 

partitioning across diff erent forests, but partitioning 

is critical to understanding allocation and requires 

additional study.

146. Luyssaert, S.; Schulze, E.-D.; Borner, A.; Knohl, 

A.; Hessenmoller, D.; Law, B.E.; Ciais, P.; Grace, J. 

2008. Old-growth forests as global carbon sinks.

Nature. 455: 213-215.

Th is publication challenges the traditional assumption 

that old-growth forests are C neutral. Data from 

519 temperate and boreal forests shows that NEP is 

positive for most forests that are 15-800 years old. 

Unmanaged primary forests constitute 30 percent of 

the global forest area; half of this area is in northern 

temperate and boreal regions. Th ese primary temperate 

and boreal forests provide 10 percent of global NEP 

and sequester 1.3 +/- 0.5 Gt C/yr, but their protection 

is not assured by current policies Release of C might be 

signifi cant if these forests are disturbed.

147. Malmsheimer, R.W.; Heff ernan, P.; Brink, 

S.; Crandall, D.; Deneke, F.; Galik, C.; Gee, 

E.; Helms, J.A.; McClure, N.; Mortimer, M.; 

Ruddell, S.; Smith, M.; Stewart, J. 2008. Forest 

management solutions for mitigating climate 

change in the United States. Journal of Forestry. 

106: 115-118.

Th is entire issue of the Journal of Forestry is devoted 

to mitigation of climate change through forest 

management, and contains the following chapter 

headings: Overview of Global Climate Change (1); 

Potential Eff ects of Climate Change on Forests 

(2); Preventing GHG Emissions through Wood 

Substitution (3); Preventing GHG Emissions 

through Biomass Substitution (4); Preventing GHG 

Emissions through Wildfi re Behavior Modifi cation 

(5); Preventing GHG Emissions through Avoided 

Land- Use Change (6); Reducing Atmospheric GHGs 

through Sequestration (7); Markets for Forest Carbon 

Off set Projects (8); and Opportunities and Challenges 

for Society, Landowners, and Foresters (Conclusion).

148. Magnani, F.; Mencuccini, M.; Borghetti, M.; 

Berbigier, P.; Berninger, F.; Delzon, S.; Grelle, A.; 

Hari, P.; Jarvis, P.G.; Kolari, P.; Kowalski, A.S.; 

Lankreijer, H.; Law, B.E.; Lindroth, A.; Loustau, 

D.; Manca, G.; Moncrieff , J.B.; Rayment, M.; 

Tedeschi, V.; Valentini, R.; Grace, J. 2007. Th e 

human footprint in the carbon cycle of temperate 

and boreal forests. Nature. 447: 848-850.

Th is comprehensive meta-analysis of northern 

hemisphere forests shows that while temporal 

disturbances and forest management play a critical role 
42

in forest C sequestration and variability, increased N 

deposition due to human activity is a primary driver 

of NEP in temperate forests. Th e high correlation 

(R2 =0.97) between average C sequestration and wet 

N deposition is often obscured by age eff ects when 

studying individual stands. Wet deposition rates are 

as high as 9.8 kg N/ha/yr, which equals about 15 kg 

N/ha/yr total N deposition. Results also show that: 

saturation is unlikely, ecosystem responses are adverse 

(including soil acidifi cation, nutrient imbalances, 

tree damage) only at deposition rates of 50-60 kg N/

ha/yr, and ecosystem response is a function of the 

addition rate rather than the cumulative N input. Th e 

C balance of temperate and boreal forests is being 

controlled and driven by human infl uence, be it direct 

(through forest management) or indirect (through N 

deposition).

149. Markewitz, D. 2006. Fossil fuel carbon 

emissions from silviculture: Impacts on net 

carbon sequestration in forests. Forest Ecology 

and Management. 236: 153-161.

Th is study highlights that few papers have quantifi ed 

and evaluated fossil emissions related to forest 

management and silvicultural activities. Th ese must 

be accounted for in order to create net positive C 

balances from forest management. Th e author reviews 

and synthesizes data ranging from peer-reviewed 

articles to equipment owner manuals to estimate C 

emissions related to silvicultural activities. Th ese data 

are used to parameterize a model for a short-rotation 

(25 year) southern pine plantation. Total C emissions 

from silvicultural activities over this period totaled 

<3 Mg C/ha. Th ese emissions are comparable to the 

amount of gains in soil C and paper product storage 

over a 100-year period for this type of stand. Th is 

study suggests that extending rotation lengths to grow 

saw-log quality trees and creating long-lasting wood 

products would “exceed the incurred C emissions 

by threefold,” that is, these practices would exceed 

emissions by 35 Mg C/ha over 100 years.

150. Montagnini, F.; Nair, P.K.R. 2004. Carbon 

sequestration: An underexploited environmental 

benefi t of agroforestry systems. Agroforestry 

Systems. 61: 281-295.

Th is publication extols the benefi ts of agroforestry 

systems, which can provide signifi cant C sequestration 

benefi ts, particularly when perennial rather 

than annual crops are planted. A review of the 

agroforestry literature is reviewed provides estimates 

of sequestration potentials in diff erent biomes. Th e 

estimated potential for enhanced sequestration 

through agroforestry in the United States is as high as 

90.3 Tg C/yr (aboveground and belowground storage). 

It includes a somewhat outdated overview of PES 

schemes in both tropical and temperate regions.

151. Myneni, R.B.; Dong, J.; Tucker, C.J.; Kaufmann, 

R.K.; Kauppi, P.E.; Liski, J.; Zhou, L.; Alexeyev, 

V.; Hughes, M.K. 2001. A large carbon sink in the 

woody biomass of Northern forests. Proceedings 

of the National Academy of Sciences. 98: 14784.

Th e authors investigate the terrestrial sink potential of 

northern temperate and boreal forests in the United 

States, Canada, and Eurasia (Finland, Norway, 

Sweden, Russia), which cover an estimated 1.4-

1.5 billion ha. Th e study examines high-resolution 

satellite data (8-km resolution, 64-km2 pixel area) 

from radiometers on board NOAA satellites and 

compares/matches them to ground-up 1980s and 

1990s inventory data from the six countries. Spatially 

explicit maps that highlight carbon pools and changes 

in C pool were developed. Results indicate C gains 

exceeding 0.3 Mg C/ha/yr in many parts of the 

Eurasian boreal and U.S. and Canadian temperate 

forests, and losses of 0.1 Mg C/ha/yr in some parts 

of the Canadian boreal forest. Th e study includes a 

table that summarizes remote sensing estimates of C 

pools and sinks broken down by country, including 

estimates for China and Japan. Average forest C pools 

are largest in the United States (57.91 Mg/ha); annual 

sinks are most signifi cant in United States (141.53 Mt 

C/yr) and Russia (283.59 Mt C/yr). Although results 

may contain inherent uncertainties and calibration 

errors, they nonetheless represent an important step 

toward understanding global C dynamics. Th is is the 

fi rst study in which high-resolution satellite estimates 

of C pool changes are validated with ground data.

152. Nabuurs, G.J.; Masera, O.; Andrasko, K.; 

Benitez-Ponce, P.; Boer, R.; Dutschke, M.; Elsiddig, 
43

E.; Ford-Robertson, J.; Frumhoff , P.; Karjalainen, 

T.; Krankina, O.; Kurz, W.A.; Matsumoto, M.; 

Oyhantcabal, W.; Ravindranath, N.H.; Sanz 

Sanchez, M.J.; Zhang, X. 2007. Forestry. In: 

Metz, B.; Davidson, O.R.; Bosch, P.R.; Dave, R.; 

Meyer, L.A., eds. Climate change 2007: mitigation. 

Contribution of working group III to the fourth 

assessment report of the Intergovernmental Panel 

on Climate Change. Cambridge, United Kingdom: 

Cambridge University Press. 851 p.

Th is chapter of the IPCC’s Working Group III 

Assessment Report, is a high-level synthesis and 

overview of the role of forests in the global C balance 

and fl ux. Th is includes an assessment of C stocks 

in living biomass by geographic regions, mitigation 

options (maintain/increase forest area; maintain/

enhance site-level C density; maintain/enhance 

landscape level C stocks; and maintain/enhance the 

use of wood products and bioenergy as substitutes for 

fossil fuel or fossil fuel intensive products), potential 

sink capacities vis-à-vis costs, adaptation strategies, and 

policy barriers and opportunities.

153. Narayan, C.; Fernandes, P.M.; van Brusselen, 

J.; Schuck, A. 2007. Potential for CO2 emissions 

mitigation in Europe through prescribed burning 

in the context of the Kyoto Protocol. Forest 

Ecology and Management. 251: 164-173.

Prescribed burning can be used to mitigate against 

catastrophic wildfi res, which result in large releases 

of CO2. Th us, prescribed burning can be considered 

a Kyoto-relevant means to reduce emissions 

countrywide. Th is study analyzes the potential for 

33 European countries to reduce emissions through 

controlled burns. Th is strategy is particularly relevant 

for wildfi re-prone countries. Results extrapolated from 

limited data indicate that on a continent-wide level, 

prescribed burning could reduce emissions by about 

50 percent. Th is reduction is based on comparing 11 

million tons CO2 released from wildfi re with 6 million 

tons released from prescribed burns; the latter strategy 

improves the likelihood that an area would not be 

burned by wildlife, but if it did burn, would release 

fewer emissions. However, the estimate in emissions 

“savings” is highly uncertain because the data set is 

limited and because it is diffi cult to quantify fuel 

consumption and emissions due to the unpredictable 

nature of wildfi res.

154. Pacala, S.W.; Hurtt, G.C.; Baker, D.; Peylin, 

P.; Houghton, R.A.; Birdsey, R.A.; Heath, 

L.S.; Sundquist, E.T.; Stallard, R.F.; Ciais, P.; 

Moorcroft, P.; Caspersen, J.P.; Shevliakova, E.; 

Moore, B.; Kohlmaier, G.; Holland, E.; Gloor, 

M.; Harmon, M.E.; Fan, S.M.; Sarmiento, J.L.; 

Goodale, C.L.; Schimel, D.; Field, C.B. 2001.

Consistent land- and atmosphere-based U.S. 

carbon sink estimates. Science. 292: 2316-2320.

It is estimated that the C sink in the Northern 

Hemisphere is 1-2 Pg per year, but the cause 

and spatial distribution of the sink are uncertain. 

Th is study clarifi es how to make consistent direct 

comparisons between land and atmospheric-based 

sinks, that is, compare the same time period, land 

area, and set of biogeochemical fl uxes. A summary 

table itemizes eight fl ux components that collectively 

create net atmosphere-to-ground fl ux, and includes 

a list and estimates from previous U.S. studies of C 

fl ux and related components. Th is study eliminates 

inconsistencies associated with constructing fl ux 

estimates for each of the eight components, and 

estimates a total sink of 0.30 to 0.58 Pg C/yr for 

1980-89. One-half of component sinks are outside the 

forestry sector.

155. Piussi, P.; Farrell, E.P. 2000. Interactions 

between society and forest ecosystems: 

challenges for the near future. Forest Ecology and 

Management. 132: 21-28.

Th is paper provides a broad overview of the ways in 

which Europeans have interacted with, infl uenced, 

and managed forests throughout history and in the 

modern era, and discusses how shifting socioeconomic 

conditions aff ect current management, including C 

outcomes. Th e second half of the paper is devoted to 

illuminating future research needs.

156. Povellato, A.; Bosello, F.; Giupponi, C. 2007. 

Cost-eff ectiveness of greenhouse gases mitigation 

measures in the European agro-forestry sector: a 
44

literature survey. Environmental Science & Policy. 

10: 474-490.

Th is paper analyzes the cost of various agricultural 

and forest-based off sets and concludes that the most 

economically viable forest-based off sets result from 

forest management. Aff orestation and bioenergy 

utilization are not as economically viable due to 

opportunity costs related to alternative land uses. 

Th ese fi ndings are similar to U.S.-based economic 

evaluations of agriculture-forestry mitigation costs.

157. Pregitzer, K.S.; Euskirchen, E.S. 2004. Carbon 

cycling and storage in world forests: biome 

patterns related to forest age. Global Change 

Biology. 10: 2052-2077.

Th is wide-reaching synthesis draws together data on 

NPP, NEP, and fi ve forest C pools (living biomass, 

CWD, organic soil horizons, soil, and total ecosystem) 

for forests in three biomes (tropical, temperate, and 

boreal) to produce information on productivity 

and variability within separate age-biome specifi c 

combinations. With minor exceptions, all C pools 

increased as stand age increased in all biomes. Carbon 

storage and cycling dynamics are closely related 

to stand age in all biomes (age plays a large role in 

variability of NEP). Additional research is needed on 

C cycle dynamics and the interaction of forest age and 

disturbance history.

158. Rice, C.W. 2006. Introduction to special 

section on greenhouse gases and carbon 

sequestration in agriculture and forestry. Journal 

of Environmental Quality. 35: 1338-1340.

Th is introduction to the “3rd USDA Symposium on 

GHG and Carbon Sequestration in Agriculture and 

Forestry,” is broad overview of the contribution and 

mitigative potential of agriculture and forestry to 

GHG management.

159. Richards, K.R.; Stokes, C. 2004. A review of 

forest carbon sequestration cost studies: A dozen 

years of research. Climatic Change. 63: 1-48.

Th is study reviews the results of more than 36 

studies on forest C economics from 1989-2001 and 

concludes that: 1) accounting inconsistency plagued 

the literature such that a ton of carbon did not always 

refer to the same quantity; 2) forest C storage capacity, 

both domestically and internationally, is substantial 

at relatively low cost (250-500 million tons in the 

United States at a cost of $10-150/ton of carbon; 

2 billion tons per year globally) for the next several 

decades; 3) there are many co-benefi ts of forest-based 

C sequestration; and 4) leakage and other accounting 

issues must be addressed or terrestrial sequestration 

may not result in real gains.

160. Van Kooten, G.C.; Eagle, A.J.; Manley, 

J.; Smolak, T. 2004. How costly are carbon 

off sets? A meta-analysis of carbon forest sinks.

Environmental Science and Policy. 7: 239-251.

Th is study provides a meta-analysis of 981 estimates 

to determine the cost of forest-based C off sets. 

Forest conservation is a relatively low-cost mitigation 

strategy, with average costs ranging from U.S. $46.62 

- $260.29/ton C. Aff orestation and agroforestry are 

relatively more expensive options among this type 

of off set. Costs generally are higher when alternative 

land-use opportunity costs are taken into account and 

marginal rather than average costs are used.

161. Volney, W.J.A.; Fleming, R.A. 2000. Climate 

change and impacts of boreal forest insects.

Agriculture, Ecosystems & Environment. 82: 283-

294.

Th e manner in which insect outbreaks and climate 

change interact with and exacerbate each other are 

described. A review of the literature associated with 

stand-replacing outbreaks in the boreal forests of 

Canada indicates that insect outbreaks can release 1.3 

- 2 times the amount of C released through wildfi res, 

and that positive feedback cycles associated with 

these outbreaks need be taken into account when 

formulating mitigation and adaptation strategies.

V. Policy Perspectives

162. Breshears, D.D.; Allen, C.D. 2002. Th e 

importance of rapid, disturbance-induced losses 

in carbon management and sequestration. Global 

Ecology & Biogeography. 11: 1-5.
45

Th ere are signifi cant risks associated with an over 

reliance on terrestrial sinks for climate change 

mitigation because the potential for rapid, disturbanceinduced losses is rarely acknowledged. Th ese risks 

become more extreme as climate change worsens, 

feedback intensifi es, and the earth’s systems approach 

potential tipping points. Th e extent to which global 

ecosystems are at risk for such rapid losses is discussed.

163. Chen, J.; Brosofske, K.D.; Noormets, A.; Crow, 

T.R.; Bresee, M.K.; Le Moine, J.M.; Euskirchen, 

E.S.; Mather, S.V.; Zheng, D. 2004. A working 

framework for quantifying carbon sequestration 

in disturbed land mosaics. Environmental 

Management. 33: S210-S221.

Th is paper provides a new framework that the authors 

recommend for future studies of net C exchange in 

disturbed land mosaics. Edge areas and related eff ects 

diff er substantially diff erent from inner landscapes and 

warrant separate consideration when the C dynamics 

of a landscape are assessed.

164. de Jong, B.H.J. 2001. Uncertainties in 

estimating the potential for carbon mitigation 

of forest management. Forest Ecology and 

Management. 154: 85-104.

Th is study quantifi es the sources of C fl ux 

uncertainties and errors and related to the classifi cation 

of land-use/land-cover types, estimation of C stocks 

within each land use/land cover, historical land use/

land cover and associated baseline assumptions, 

and variations in parameter values used to calculate 

carbon fl uxes. Inclusion of forestry into the CDM is 

recommended.

165. Depro, B.M.; Murray, B.C.; Alig, R.J.; Shanks, 

A. 2008. Public land, timber harvests, and climate 

mitigation: Quantifying carbon sequestration 

potential on U.S. public timberlands. Forest 

Ecology and Management. 255: 1122-1134.

Th is study provides a reference and basis for 

comparison for policymakers and forest managers 

considering C sequestration and associated economic 

potential of diff erent harvest scenarios in the United 

States. Th ree simplifi ed scenarios are considered: 

business-as-usual, complete elimination of harvesting, 

and a return to pre-1989 harvesting levels. Th e latter 

two scenarios resulted in increases and decreases, 

respectively, in forest C storage on the order of 50 

percent. However, the study does not consider how 

the risk of wildfi re or other disturbances aff ects these 

scenarios, or the extent to which calculations might be 

altered by incorporating biofuels. Th e economic value 

of sequestered C on public forests could exceed timber 

values should these forests participate in C markets.

166. Im, E.H.; Adams, D.M.; Latta, G.S. 2007. 

Potential impacts of carbon taxes on carbon fl ux 

in western Oregon private forests. Forest Policy 

and Economics. 9: 1006-1017.

Th e eff ects of accounting for C revenues and costs 

in forest management decisions on both industrial 

and NIPF lands in western Oregon are considered. 

Results show that C taxes increase rotation lengths 

and concentrate production on lands with higher 

productivity. Taxes also reduce consumer and 

producer surpluses, though producer surplus losses 

are partially off set by C revenues. Th e baseline and tax 

rate used strongly infl uences outcomes and landowner 

participation. Taxes could be a useful policy tool 

to spur greater terrestrial sequestration, though 

disturbance and leakage risks must be taken into 

account. Th ese risks likely will increase as a result of 

increasing land-based C stocks.

167. Kirschbaum, M.U.F. 2003. To sink or burn?

A discussion of the potential contributions 

of forests to greenhouse gas balances through 

storing carbon or providing biofuels. Biomass and 

Bioenergy. 24: 297-310.

Th is study emphasizes that although forests play 

an important role in the global C cycle, land-based 

sinks are subject to saturation and land management 

alone cannot mitigate the buildup of atmospheric 

CO2 concentrations. Storage in wood products and 

displacement of fossil fuels through bioenergy use can 

have relatively small but meaningful impacts on overall 

atmospheric C concentration in the near-term; over 

time, substituting bioenergy for fossil fuel results in 

signifi cant benefi ts.
46

168. McCarl, B.A.; Schneider, U.A. 2001. Climate 

change: Greenhouse gas mitigation in U.S. 

agriculture and forestry. Science. 294: 2481-2482.

Th e role of agriculture and forestry mitigation is 

delineated in three main categories: reduction of direct 

emissions; expansion of the terrestrial carbon sink; and 

production of products that substitute for fossil-fuel 

intensive ones. However, the model used (Agricultural 

Sector Model, including GHG; ASMGHG) focuses 

almost exclusively on management alternatives, 

tradeoff s, and economic impacts for agriculture, 

and does not explicitly address forest management 

activities, an important factor in C sink expansion.

169. Nabuurs, G.J.; Th ürig, E.; Heidema, N.; 

Armolaitis, K.; Biber, P.; Cienciala, E.; Kaufmann, 

E.; Mäkipää, R.; Nilsen, P.; Petritsch, R.; Pristova, 

T.; Rock, J.; Schelhaas, M.J.; Sievanen, R.; 

Somogyi, Z.; Vallet, P. 2008. Hotspots of the 

European forests carbon cycle. Forest Ecology and 

Management. 256: 194-200.

Article 3.3 of the Kyoto Protocol mandates 

accounting and reporting with respect to aff orestation, 

reforestation, and deforestation, but article 3.4 allows 

Annex I countries to decide whether to include forest 

management activities that increase sink strength and 

use these activities to partially meet GHG reduction 

requirements. Of 25 European countries, 17 elected 

to include forest management activities. Th is paper 

summarizes how certain forest management activities 

infl uence C pools and fl uxes over various time scales 

and emphasizes the importance of local solutions 

and appropriate management that integrates goals 

and optimizes outcomes based on regional and local 

factors.

170. Noss, R.F. 2001. Beyond Kyoto: Forest 

management in a time of rapid climate change. 

Conservation Biology. 15: 578-590.

Th is study suggests forest management practices that 

can help create and restore ecosystem and biodiversity 

resilience in an era of rapid and unpredictable change. 

Key among these are protecting primary forests, 

maintaining and creating connectivity, creating reserve 

buff er zones, practicing low-intensity forestry and 

maintaining natural forests (avoiding conversion to 

plantations), and preserving and maintaining keystone 

species, diverse gene pools, and natural fi re patterns.

171. Pacala, S.; Socolow, R. 2004. Stabilization 

wedges: Solving the climate problem for the next 

50 years with current technologies. Science. 305: 

968-972.

Th is paper describes a toolbox approach to solving 

the climate crisis that includes 15 approaches, each 

of which would reduce emissions by 1 Gt C/yr. To 

reach climate stabilization by 2054, society would 

need to adopt and implement a total of seven wedges. 

Two stabilization wedges are related to agriculture 

and forestry: conservation tillage applied to cropland 

globally and halting tropical deforestation along with 

establishing 300 million ha of new tree plantations. 

Each wedge could reduce emissions by 1Gt C/yr.

ACKNOWLEDGMENTS

We acknowledge fi nancial support from the USDA 

Resources Planning Act Assessment Program. We 

also appreciate the review and comments from three 

anonymous reviewers and Maria Janowiak, Michigan 

Technological University. Eric Fiegenbaum provided 

graphics for the cover.
47

APPENDIX

Unit of measure Explanation

Mg (Megagram) = 106 grams = 1 metric ton = 1 tonne

Mt (Megaton) =106 tons = 1,000,000 tonnes

Tg (Teragram) = 1012 grams = 1 million metric tons = 1,000,000 tonnes

Pg (Petagram) = 1015 grams = 1 billion metric tons = 1,000,000,000 tonnes

ha (hectare) = 2.471 acres

Mha = million hectares

ac (acre) = 0.405 hectares

Table A1.—Units of measures and conversions

Abbreviation Name

CO2 Carbon dioxide

CO4 Carbon tetraoxide

NO2 Nitrous dioxide

N2O Nitrous oxide

N Nitrogen

P Phosphorous

K Potassium

Table A2.—Names of chemical elements 

and compounds and their abbreviation
48

Table A3.—Acronym or abbreviations and their defi nitions

Abbreviation/

acronym

Term

ANPP Aboveground net primary productivity

BT Basal thinning

BOREAS Boreal Ecosystem-Atmosphere Study (Canada)

CDM Clean development mechanism

CERs Certifi ed emission reductions

CO2e Carbon dioxide equivalent; the amount of global warming potential generated, using 

carbon dioxide as a reference

CWD Coarse woody debris

DOM Dead organic matter

FACE Free-air CO2 enrichment

FFC Forest fl oor carbon

FIA Forest Inventory and Analysis, the name of a program within the U.S. Forest Service

FWD Fine woody debris

GHG Greenhouse gas

GPP Gross primary productivity

IPCC Intergovernmental Panel on Climate Change

MAI Mean annual biomass increment

MCA Multi-criteria analysis

NDVI Normalized difference vegetation index

NEP Net ecosystem productivity

NIPF Nonindustrial private forest

NOAA National Oceanographic and Atmospheric Administration

NPP Net primary productivity

PES Payment for ecosystem services

ppm Parts per million

REDD Reduced emissions from deforestation and degradation

RMS Reclaimed mine soil

SEV Soil expectation value

SOC Soil organic carbon

SOM Soil organic matter

SRR Soil respiration response

TBCA Total belowground carbon allocation

UNFCCC United Nations Framework Convention on Climate Change
49

Table A4.—Scientifi c and common names of tree species referenced in the studies

Latin Name Common Name

Abies alba silver fi r

Abies balsamea balsam fi r

Abies fraseri Fraser fi r

Acer spp. maple

Betula alba white birch

Betula pendula silver birch, also known as European white birch

Carya cordiformis bitternut hickory

Carya glabra pignut hickory

Chamaecyparis obtuse Hinoki falsecypress

Crytomeria japonica Sugi (cedar)

Eucalyptus regnans (Australian) mountain ash/swamp gum

Fagus spp. beech

Fagus sylvatica European/common beech

Fraxinus spp. ash

Larix kaempferi Japanese larch

Picea glauca white spruce

Picea mariana black spruce

Picea rubens red spruce

Picea sitchensis Sitka spruce

Pinus banksiana jack pine

Pinus contorta lodgepole pine

Pinus elliotti slash pine

Pinus pinaster maritime pine

Pinus ponderosa Ponderosa pine

Pinus radiata radiate pine

Pinus resinosa red pine

Pinus rigida pitch pine

Pinus sylvestris Scots pine

Pinus taeda loblolly pine

Platanus occidentalis American sycamore

Populus deltoides eastern cottonwood

Populus tremuloides trembling aspen

Populus trichocarpa black cottonwood

Pseudotsuga menziesii Douglas-fi r

Quercus spp. oak 

Quercus alba white oak

Quercus coccinea scarlet oak

Quercus robur English oak

Quercus velutina black oak

Tilia spp. basswood

Tsuga heterophylla western hemlock
The U.S. Department of Agriculture (USDA) prohibits discrimination in all its programs and activities on the basis of 

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call (800)795-3272 (voice) or (202)720-6382 (TDD). USDA is an equal opportunity provider and employer. 
Hines, Sarah J.; Heath, Linda S.; Birdsey, Richard A. 2010. An annotated 

bibliography of scientifi c literature on managing forests for carbon benefi ts.

Gen. Tech. Rep. NRS-57. Newtown Square, PA: U.S. Department of Agriculture, 

Forest Service, Northern Research Station. 49 p. [Includes CD-ROM].

Managing forests for carbon benefi ts is a consideration for climate change, bioenergy, 

sustainability, and ecosystem services. A rapidly growing body of scientifi c literature on 

forest carbon management includes experimental, modeling, and synthesis approaches, 

at the stand- to landscape- to continental-level. We conducted a search of the scientifi c 

literature on the topic of managing forests for carbon, and compiled an annotated 

list of citations. We chose to focus specifi cally on studies that addressed carbon in 

aboveground carbon pools, at both the micro (tree, stand) and macro (landscape, 

policy) levels. Aboveground pools include: live tree, understory, standing dead wood, 

down dead wood, and forest fl oor. The temporal scope of the literature search was the 

period 2000-2008 and the geographical scope was the temperate and boreal forests 

mainly in the United States, but also Canada, Europe, Russia, Japan, China, New 

Zealand, and Australia.

KEY WORDS: forest management, carbon sequestration, mitigation, climate change

The contents of the cd are available 

on the NRS Publications web page: 

http://www.nrs.fs.fed.us/pubs/34886
www.nrs.fs.fed.us