THE HARVARD FOREST 2001-2002

Harvard University

Front Cover: Walk-up towers erected in numerous locations throughout theForest provide access for studies at different levels in the forestcanopy as well as panoramic views across the landscape.

Back Cover: The Prospect Hill fire tower as seen from the hemlock standwalk-up tower.

Photography by David Foster, John Burk and, Amanda Brown.

Contents

Personnel at the Harvard Forest - - - - - 4 Introduction to the Harvard Forest - - - - - 5 New Staff - - - - - - - - 6Transitions and Accolades - - - - - - 6Research Activities - - - - - - - 7Bullard Fellows - - - - - - - - 22Educational Activities - - - - - - - 24

Summer Research Program - - - - - 24Activities of the Fisher Museum - - - - - 24

Meetings, Conferences, Seminars - - - - 26Forest Management and Maintenance - - - - - 26Computers - - - - - - - - 27Activities of the Harvard Forest Staff - - - - - 28Visiting Research Scientists - - - - - - 30Publications - - - - - - - - 31Acknowledgment of Support and Gifts - - - - - 32Gifts - - - - - - - - - 33New Funding - - - - - - - - 33

ANNUAL REPORT OF THE HARVARD FOREST 2001-2002

http://harvardforest.fas.harvard.edu

Audrey Barker Plotkin Research AssistantSylvia Barry Musielewicz Research AssistantJesse Bellemare Graduate StudentAllison Berry Bullard FellowEmery Boose Information and

Computer System Manager

Jeannette Bowlen AccountantJohn Burk Archivist Tawana Childs Laboratory TechnicianRichard Cobb Research AssistantElizabeth Colburn Bullard FellowWillard Cole Woods CrewElaine Doughty Research AssistantJohn Edwards Forest ManagerEdythe Ellin AdministratorAaron Ellison Bullard Fellow and

Senior EcologistAdrian Fabos Facilities ManagerSamantha Farrell Laboratory TechnicianBarbara Flye Librarian/SecretaryCharles H. W. Foster AssociateDavid Foster DirectorDonna Francis Research AssociateKelli Graves Secretarial AssistantLucas Griffith Woods CrewJulian Hadley EcophysiologistBrian Hall Research AssistantJulie Hall Research Assistant Linda Hampson Secretarial Assistant

Teresa Jones Assistant Schoolyard Coordinator

David Kittredge Forest Policy AnalystMatt Kizlinski Graduate StudentOscar Lacwasan Woods CrewDana MacDonald Research AssistantLisa Marselle Summer CookGlenn Motzkin Plant EcologistJohn O’Keefe Museum and

Schoolyard Coordinator

Colin Orians Bullard FellowDavid Orwig Forest EcologistJulie Pallant System and Web

AdministratorSarah Parnes Graduate StudentTim Parshall Post-doctoral FellowAkile Pite Bullard FellowDorothy Recos-Smith Staff AssistantJessica Schedlbauer Research AssistantLaura Schreeg Summer Program

AssistantMichael Scott Woods CrewKimberly Smith Bullard FellowNavjot Sodhi Bullard FellowMindy Syfert Research AssistantP. Barry Tomlinson E. C. Jeffrey Professor of

Biology, EmeritusJohn Wisnewski Woods Crew

Steven Wofsy Associate

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PERSONNEL AT THE HARVARD FOREST 2001-2002

INTRODUCTION TO THE HARVARD FOREST

Since its establishment in 1907, the Harvard Foresthas served as a center for research and education inforest biology. Through the years researchers havefocused on silviculture and forest management, soilsand the development of forest site concepts, the biol-ogy of temperate and tropical trees, forest ecology,forest economics, landscape history, conservationbiology, and ecosystem dynamics. Today, this legacyof research and education continues as faculty, staff,and students seek to understand historical and mod-ern changes in the forests of New England andbeyond resulting from human and natural distur-bance processes, and to apply this information to theconservation, management, and appreciation of natu-ral ecosystems. This activity is epitomized by theHarvard Forest Long Term Ecological Research (HFLTER) program, which was established in 1988through funding by the National Science Foundation(NSF).

Physically, the Harvard Forest is comprised ofapproximately 3,000 acres of land in the north-cen-tral Massachusetts town of Petersham, which includemixed hardwood and conifer forests, ponds, exten-sive spruce and maple swamps, fields, and diverseplantations. Additional land holdings include the 25-acre Pisgah Forest in southwestern New Hampshire(located in the 5000-acre Pisgah State Park), a virginforest of white pine and hemlock that was 300 yearsold when it blew down in the 1938 Hurricane; the100-acre Matthews Plantation in Hamilton,Massachusetts, which is largely comprised of planta-tions and upland forest; and the 90-acre Tall TimbersForest in Royalston, Massachusetts. In Petersham acomplex of buildings that includes Shaler Hall, theFisher Museum, and the John G. Torrey Labora-tories provide office and laboratory space, computerand greenhouse facilities, and a lecture room for sem-inars and conferences. Nine additional houses pro-vide accommodations for staff, visiting researchers,and students. Extensive records, including long-termdata sets, historical information, original field notes,maps, photographic collections, and electronic dataare maintained in the Harvard Forest Archives.

Administratively, the Harvard Forest is a depart-ment of the Faculty of Arts and Sciences (FAS) ofHarvard University. The Harvard Forest administersthe Graduate Program in Forestry that awards a mas-ter’s degree in Forest Science and faculty at the Forestoffer courses through the Department of Organismic

and Evolutionary Biology (OEB), the KennedySchool of Government (KSG), and the FreshmanSeminar Program. Close association is also main-tained with the Department of Earth and PlanetarySciences (EPS), the School of Public Health (SPH),and the Graduate School of Design (GSD) atHarvard and with the Department of NaturalResource Conservation at the University of Massa-chusetts, the Ecosystems Center of the MarineBiological Laboratory at Woods Hole, and theComplex Systems Research Center at the Universityof New Hampshire.

The staff and visiting faculty of approximatelyfifty work collaboratively to achieve the research,educational, and management objectives of theHarvard Forest. A management group meets monthly to discuss current activities and to planfuture programs. Regular meetings with the HFLTER science team, weekly research seminars andlab discussions, and an annual ecology symposiumprovide for an infusion of outside perspectives. Thefive-member Woods Crew and Forest Managerundertake forest management and physical plantactivities. The Coordinator of the Fisher Museumoversees many educational and outreach programs.

Funding for the Harvard Forest is derived fromendowments and FAS, whereas major research sup-port comes primarily from the National ScienceFoundation, Department of Energy (NationalInstitute for Global Environmental Change), U.S.Department of Agriculture, NASA, Andrew W.Mellon Foundation, and other granting sources. Oursummer Program for Student Research is supportedby the National Science Foundation, the Andrew W.Mellon Foundation, and the R. T. Fisher Fund.

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NEW STAFF

Aaron Ellison, a recent Bullard Fellow and MarjorieFisher Professor of Environmental Studies at Mt.Holyoke College, accepted a position as SeniorEcologist at Harvard Forest. Aaron, who received hisB.A. from Yale University and Ph.D. from BrownUniversity, will continue his research in population,community, and statistical ecology.

Adrian Fabos, a graduate of Carnegie MellonUniversity and the Yale School of Forestry, replacedJohn Edwards as Facilities Manager and becameimmediately involved in supervising the WoodsCrew, contractors, and facilities operations.

Sarah Parnes arrived this summer to begin herstudies in the Master’s in Forestry Science program.Sarah, a recent graduate of the University of Virginiawith a B.A. in environmental science, will be workingclosely with David Foster and Glenn Motzkin inresearching the history of white pine and hemlockforests in Petersham.

Julie Hall, a new research assistant, is workingclosely with Dave Orwig, Glenn Motzkin, and BrianHall on a series of remote sensing and (GeographicInformation System (GIS) projects. Lucas Griffithand Michael Scott joined the Woods Crew, replacingretirees Donald Hesselton and Pete Spooner.

TRANSITIONS AND ACCOLADES

On July 1, 2001, Barry Tomlinson retired with thetitle “E. C. Jeffrey Professor of Biology Emeritus,” butcontinued his research at Harvard Forest. This repre-sents a continuation of an association with HarvardForest that began with a visit to Petersham inDecember 1961. Concomitant with his retirementBarry was appointed Eleanor Crum Professor ofTropical Botany at the National Tropical BotanicalGarden, Kalaheo, Hawaii and will divide his timebetween the Forest and Hawaii. In April 2002 the

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Sarah Parnes.

Michael Scott.

Aaron Ellison.

Smithsonian Institution, Washington, D.C., awardedBarry Tomlinson the José Cuatracasas Medal at itsconvention on Biological Diversity. The medal isawarded for Excellence in Tropical Botany.

John “Jack” Edwards retired after eighteen yearsas Facilities Manager and Forest Manager at theHarvard Forest, where he dedicated himself toimproving the working and living conditions for allstaff and visitors. Just a few of his accomplishmentsinclude: overseeing the complete renovation of FisherHouse into housing for visiting researchers and sum-mer students; construction of the Pole Barn and recy-cling shed; renovation of the garage into modernarchival space and the Torrey Laboratory into amulti-user facility; erection of the EnvironmentalMonitoring Station (EMS) one mile into the woods;major landscaping projects around and significantrenovation projects within Shaler Hall; and structuralimprovements to every residence and buildingowned by the Forest. Jack’s impact on the Forest willbe evident for many years to come.

RESEARCH ACTIVITIES

An Historical Approach to Understanding and Conserving the Coastal Region

Following up on our previous studies in centralMassachusetts, the Connecticut Valley, and Martha’sVineyard, we are conducting a comprehensive inves-tigation of the history and vegetation of the coastalregion that includes Cape Cod, Martha’s Vineyard,Nantucket, Block Island, and Long Island. Thisregion is a high priority for conservation because itsupports numerous rare or uncommon plant and ani-mal species and communities and is highly threat-ened by development. With support from theNational Science Foundation, Andrew MellonFoundation, Massachusetts Biodiversity Initiative,and The Nature Conservancy’s Ecological ResearchProgram, we are investigating the link between thelandscape history of the coastal region and the mod-ern abundance and distribution of upland plant com-munities, including grasslands, heathlands, barrens,and woodlands. Although the region has a long his-tory of settlement and intensive use, there has beenno prior attempt to rigorously evaluate the impact of

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Barry Tomlinson.

John Edwards.

historical land use on species distributions, commu-nity assemblages, or biodiversity in this region. Suchan understanding is critical in order to understandmodern vegetation patterns, to identify appropriategoals for conservation, and to develop managementapproaches for achieving those objectives.

Current work emphasizes integrated paleoeco-logical, historical, (GIS), and field studies throughoutthe coastal region. Tim Parshall and David Foster ledpaleoecological studies to reconstruct the long-termvegetation and disturbance history of the region, andBrian Hall is gathering a wide range of historicaldata, including excellent historical maps that arebeing incorporated into a growing coastal GIS database. Julie Hall developed maps of the coastal regionfrom the 1930s, enabling us to evaluate the timingand rate of reforestation after abandonment of agri-cultural lands. Dana MacDonald and Rob Eberhardtworked closely with David and Glenn Motzkin toevaluate the relationship between modern vegetationvariation, site factors, and disturbance history.

Results to date suggest that despite considerablevariation in environmental conditions and fire, thepattern and intensity of historical land-use activitiescontinue to influence modern species distributions,even many decades after these activities have ceased.In particular, several species are slow to recolonizeformer agricultural lands even in cases where mod-ern site conditions appear suitable for these species.

Forest Composition and Distribution in the HistoricalPeriod: Providing the Context for Ecological andBiodiversity Assessments for Massachusetts

Because the Massachusetts landscape has a long his-tory of human settlement, ecological study, conserva-tion, and long-term planning must incorporate anunderstanding of the history of human landscapemodification. However, to date, most ecological andplanning studies have failed to evaluate this historyeffectively, primarily due to a lack of information.With support from the Massachusetts NaturalHeritage and Endangered Species Program, DavidFoster, Glenn Motzkin, Mindy Syfert, and Brian Halldeveloped GIS data layers to assess land-use historythroughout the Commonwealth. Using maps gener-ated for each town in Massachusetts in 1830, wedeveloped maps depicting the historical extent of for-est land, open land, meadows, and cultural features,including roads, mills, meeting houses, etc. Because1830 was near the period of maximum agriculturalland clearance for many towns in the Common-wealth, the resulting data layers will provide a rea-sonable approximation of the Massachusetts land-scape during the agricultural period. Such informa-tion is valuable for a wide range of research, plan-ning, and management efforts, including establishingobjectives for conservation, restoration, long-termecological management, and archaeological and cul-tural resource assessment. Interestingly, our results

8

Tim Parshall.

Brian Hall.

indicate that the greatest extent of forest land inMassachusetts in 1830 occurred in southeasternMassachusetts, where large portions of Plymouthand Bristol counties and the inner portion of CapeCod remained forested. In contrast with the modernlandscape, the uplands of central and westernMassachusetts supported numerous small and iso-lated woodlands in a predominantly agriculturallandscape.

John Burk worked with David, Glenn, and Brianto gather data on forest composition in southern NewEngland at the time of European settlement. In com-bination, these historical sources have greatlyincreased our understanding of forest composition insouthern New England in the early historical periodand the changes that resulted from intensive agricul-tural clearing through the mid-nineteenth century.We also collaborated with Charlie Cogbill from theHubbard Brook research program, who has similardata for northern New England to evaluate early his-torical forest composition across all of New England.

Historical Land Cover and Land-use Patterns

Much past research at Harvard Forest has investigat-ed the links between historical land use and modernvegetation characteristics at both local and regionalscales. Local-scale studies have shown that there is astrong link between historical land-use practices andmodern species composition. For example, cultiva-tion eradicates existing woodland vegetation, causingformally plowed fields to lack species that are slow to recolonize, even though these species may beabundant in adjacent woodlots. Regional studieshave shown that historical land-use practices havereduced much of the climatic control over forestspecies composition and reduced regional variationin composition. In order to evaluate the relative influ-ence of climate and land use across a region with agreater environmental gradient, Brian Hall, Glenn

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John Burk.

Team Paleo preparing to retrieve asediment core from Lily Pond, NewSalem, Massachusetts. From left: ElaineDoughty, Sylvia Barry Musielewicz,Kate Musgrove, Gina Yazzie, andSarah Parnes.

Motzkin, and David Foster are investigating howforests in Massachusetts have changed over morethan 300 years of intensive land use. Preliminaryresults suggest that even though most of the state’sforests have at one time or another been cleared andused as cropland or pastureland, climate continues toinfluence species composition, although land-use hasmodified the relative abundance of tree taxa. Theintroduction of non-native pest species within the lasteighty years has also had, and will continue to have,a great effect on forest composition.

As part of the coastal project, colonial records forLong Island at the New York State Archives wereresearched for witness tree and agricultural land-usecitations. Data sets of fifty or more trees were col-lected for six towns, including Hempstead andOyster Bay in Nassau County. The year range ofthese records was from the mid-1600s to the 1720s.To help interpret the land-use history of theHempstead Plain, agricultural resolutions in thetown’s records from 1655 to 1845 were collated andsummarized. During this process, as part of a collab-oration with visiting researchers from Belgium, earlyDutch records from the Albany area including mapsand proprietor data were added to the Archives.

Hemlock Woolly Adelgid Impacts on New EnglandForests: Stand Dynamics, Ecosystem FunctionChanges and Landscape Patterns

A large research effort led by Dave Orwig is investi-gating the impact of the introduced insect, the hem-lock woolly adelgid (HWA) on forests across southernNew England. HWA is spreading from southernConnecticut, where it killed many thousands of acresof forest, and it has the potential to eliminate easternhemlock across its now native range. Vegetationdynamics have been monitored for six years in eighthemlock stands with varying levels of HWA damagein Connecticut to examine patterns of hemlock mor-tality and subsequent community reorganization fol-lowing infestation. Since 1995, overstory and under-story mortality has risen 5% to 15% per year to over-all values of 50% to 99%. There has been no treerecovery and the health of remaining trees deteriorat-ed in all infested stands, although on some sites hem-locks can remain alive for over ten years following ini-tial infestation and remain standing for six to eightyears following mortality. Rapid establishment andgrowth by black birch occurred at most sites alongwith lesser amounts of red maple and oak species.

The few hemlock seedlings found contained HWAinfestation. Shrub cover remains low following infes-tation while herbaceous cover, consisting primarily ofhay-scented fern, gradually increases over time. Tree-ring analysis of hardwood and declining hemlocktrees, coupled with age-structure analysis of newlyestablished birch saplings was effective in determiningthe timing of initial HWA impact in stands ofunknown infestation date. Over the last ten years,radial growth of hemlock declined precipitously inmost stands, while oak and maple growth exhibitedconcomitant, large annual increases (see figure page12). Results suggest that dramatic stand structure andcomposition changes accompany heavy HWA infesta-tions that continue long after hemlock mortality.

During the summer of 2002, Dave Orwig,Richard Cobb, and summer student Brandon Burkeresampled permanent plots established in 1997 toexamine the effect of hemlock decline and mortalityon nitrogen cycling. Soil analyses including pH, tem-perature, carbon:nitrogen ratios, texture, and totalsoil organic matter have been completed and

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Impacts of the hemlock woolly adelgid.

nitrogen mineralization continues to be measured ina subset of sites. Stands with heavy HWA infestationhad lower surface soil moisture levels, higher soiltemperatures and light, and higher net nitrogen min-eralization and net nitrification rates than uninfestedstands or stands with low damage. Resin bags buriedin the soil at damaged sites also captured higheramounts of ammonium and nitrate than similar bagsin uninfested stands, indicating that nitrogen isbecoming more available as a result of the microen-vironmental changes associated with HWA damage.We will continue to sample these stands to examinethe long-term changes in N cycling associated withhemlock deterioration and eventual replacement byhardwoods.

Dave, Richard Cobb, Audrey Barker-Plotkin andsummer student, Jacqueline Guzman established astudy on vegetation, soil nitrogen cycling, andmicroenvironments in two hemlock forests onHarvard Forest that lack HWA to collect baselineinformation in anticipation of the imminent arrival ofthe adelgid. Although HWA occurs elsewhere inPetersham it has only been found on one tree at theHarvard Forest. In a related effort, Audrey BarkerPlotkin and summer student Jackie Guzman gathereddetailed baseline information for several hemlockstands that have well-documented, long-term vegeta-tion histories reconstructed via pollen and tree-ring

analyses. Audrey and Jackie are rating hemlock treevigor, surveying regeneration of hemlock and othertrees, and compiling species lists of understory vege-tation. In these healthy, intact stands, understoryplants are sparse, and comprise a suite of shade-tolerant herbs, ferns, and tree seedlings. They arealso measuring ecosystem parameters such as organ-ic matter depth and nitrogen cycling. Some of thestands have a remarkably thick organic layer. Thesemeasurements will link the long-term history of thesehemlock stands to continuing studies of hemlockstand response to adelgid infestation. The proximityof these sites to Harvard Forest research facilities and scientists makes them excellent areas to expandresearch on the forest ecosystem impacts of this inva-sive pest.

Under the direction of Richard Cobb, we con-tinue to compare the rates of decomposition at thesoil surface versus the organic-mineral soil interfaceat twelve sites throughout Connecticut and Massa-chusetts. We have also begun a reciprocal litter studyat a subset of these sites comparing hemlock, blackbirch, and hemlock-black birch mixed litter decom-position. Black birch is the most prolific tree speciesestablishing on former hemlock stands. The foliarchemistry of black birch is substantially differentfrom that of hemlock and decomposition will mostlikely be different as well.

As a follow-up to our large-scale project examin-ing HWA distribution in Connecticut, Dave Orwigand summer students Matthew Burr and NicholasPovak have begun sampling hemlock stands inMassachusetts. They are mapping the distribution ofhemlock stands > 3 ha in a region encompassing theConnecticut River Valley and stretching from West

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David Orwig.

Audrey Barker Plotkin.

Springfield to Wales in the south and from Colrain toRoyalston in the north. They will obtain informationon forest structure and composition, crown vigor, sitecharacteristics, presence of HWA, and the extent andspatial patterns of damage generated by this insectsince the time of its arrival in Massachusetts in 1989.This information will be incorporated into a GISanalysis of landscape-level, biological, edaphic, andhistorical factors that control the damage patternsobserved in hemlock.

Carbon Exchange of Temperate Forests

Julian Hadley continued to measure the carbon bal-ance of various forest types on the Prospect Hill tractto develop better carbon exchange models and moreaccurate estimates of carbon balance for central NewEngland forests. New England’s forests affect globalcarbon exchange, atmospheric CO2 concentration,and the greenhouse effect resulting from globally ris-ing atmospheric CO2.

During 2000 and early 2001 work in the old-growth hemlock stand near the Black Gum Swamp

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Overstory radial growth patterns in hemlock and oak trees and the patterns of black birch establishment during chronic hemlock woolly adelgid infestations at the Foster’s Pond and Burnham Brook hemlock sites

in southern Connecticut. N = number of trees used to construct the average growth curve. As hemlock trees deteriorate, hardwood tree growth exhibits dramatic increases and dense birch seedlings rapidly become

established.

measured carbon exchange above the canopy, as wellas soil respiration. The hemlock forest stored 3.0Mg/ha during the year from November 2000 throughOctober 2001, based on data collected during periods of suitable wind direction and turbulence forcarbon exchange measurements. Carbon exchangemodels based on these measurements were used toestimate storage during other periods. Annual car-bon storage of 3.0 Mg/ha is high relative to estimatesfrom a carbon exchange model and it is much greaterthan annual above-ground carbon storage in woodduring the 1990s (1.0 to 1.4 Mg/ha). However, meas-urements by Steve Wofsy at the Harvard Forest’sEnfironmental Monitoring Station (EMS) tower indi-cated that November 2000 through October 2001was a year of record high annual carbon uptake;these environmental conditions may have alsoenhanced carbon storage in the hemlock forest.

The highest monthly estimates of net carbonstorage in the hemlock forest occurred during Apriland May 2001, while slight losses of carbon were esti-mated for the winter months and August 2001 (seefigure page 14). These observations support othermeasurements and indicate that when soil and airtemperatures were very high, total respiration of theforest could exceed photosynthesis, leading to carbonloss. August 2001 was the hottest month of the year,with monthly average soil and air temperatures bothabout 2ºC hotter than in any other month. Theannual pattern suggests that mild spring weatherenhances carbon uptake by hemlock forests, but hotsummers reduce it.

Carbon exchange measurements in a new area,the west slope of Little Prospect Hill, began in May2002. This area is dominated by red oak, red maple,and birch, with a red pine plantation about 250 mSSW of the measurement tower. The area withinabout 200 m of the tower to the SW, and 300 m tothe NW, is covered by forest only about forty-fiveyears old, originating after a severe fire in 1957.These upper slopes are also very dry. Carbonexchange data from this site will provide a valuablecontrast to the hemlock forest and the area aroundthe EMS tower, which has forest of similar speciescomposition 70 to 100 years old. The EMS tower isalso in a lower slope position where water stress isless likely, particularly to the NW where there is alarge bog. The Little Prospect Hill site will provide anopportunity to test the general applicability of theobservation from the EMS tower that drought resultsin higher than average carbon storage, because soil

respiration is reduced more than photosynthesis.This may not hold true at the new drier site.

Long-term Forest Measurements and HistoricalCarbon Sequestration

Permanent plots provide an empirical understandingof forest change over time, and are an invaluable partof forestry and ecological research. Such studies pro-vide information on stand development and canaddress numerous ecological questions, includingmany which could not have been anticipated whenthe study was initiated. Walter Lyford began meas-urements of a seven-acre area of red oak-red mapleforest on the Prospect Hill tract of Harvard Forest in 1969 when he mapped all trees greater than twoinches diameter at a very large-scale (1 inch = 5 feet).The maps include live and dead trees, stumps, wind-throws and other features such as stonewalls, boulders, soil moisture, and a damage boundaryfrom the 1938 hurricane. Last summer, AudreyBarker Plotkin and summer student Kristin Wilsonrelocated and remeasured all trees, completing the

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Detail of one of Walter Lyford’s original maps showingtree species and sizes, stumps, windthrown trees, boul-ders and a boundary between forest severely damagedby the 1938 hurricane (above dashed line) and lightlydamaged areas (below dashed line). Each grid square

represents 5 x 5 feet on the ground.

fourth measurement in the site’s thirty-two-year his-tory. They also digitized all of Lyford’s original mapson GIS.

The role of temperate forests as a major carbonsink is currently a focus of much research at HarvardForest, centered on the Environmental MonitoringStation run by Steve Wofsy in the Department ofEarth and Planetary Sciences. The Lyford data canbe used to provide an independent estimate of car-bon sequestration as one stand ages from 70 and 100years. Carbon stored in the above-ground live woodybiomass at the Lyford Grid area rose from 71 metrictons of carbon per hectare in 1969 to 114 tons perhectare in 2001, mostly in red oak. Over the 32-yearperiod, an average of 1.3 tons of carbon per hectarewere stored each year in live woody biomass.Additional pools of carbon include the soil, deadwood and roots.

Kathleen Donohue of the Department of Organismicand Evolutionary Biology conducted several studieson the evolutionary ecology and genetics of naturalplant populations. The first project investigates thegenetic basis and adaptive significance of germinationresponses to seasonal cues in Arabidopsis thaliana. Thefield component of this large quantitative geneticsexperiment has been completed, and results present-ed at the meetings of the American Society ofNaturalists. In addition, two summer ResearchExperience for Undergraduates (REU) students,Joseph Seggio and Naomi Wender, are conductingindependent projects to investigate the genetic basisof germination in Arabidopsis. Another major projectaddresses the phenotypic and genetic basis of seeddispersal in Arabidopsis thaliana. The data collectionfor this study is nearing completion, and the summer

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Monthly carbon exchange in the old-growth hemlock forest fromNovember 2000 through October 2001. Positive numbers indicate carbon

uptake by the forest, negative numbers show carbon lost to theatmosphere.

Population Ecology

REU students and Kathleen are participating in this endeavor. The third project concerns seed dis-persal in a group of mustards that have evolved to inhabit marine strands. A trip to the EuropeanSynchrotron Radiation Facility provided high resolu-tion 3-dimensional X-ray images of fruit morphology,which revealed structural features that may have con-tributed to the ability of these species to inhabitmarine habitats and to expand their ranges globally.

Kristina Stinson’s research is centered mainly onthe population biology and ecophysiology of theinvasive plant species, garlic mustard (Alliaria peti-olota) and its spread in and around Harvard Forest,as part of LTER investigations on invasive species.She is investigating the role of source-sink dynamicsin the invasion process and possible managementimplications. Seeds will be collected from garlic mus-tard populations from southern New Jersey toOntario with the objective of including a latitudinaltransect in ongoing comparisons of different light andmoisture habitats. In addition, similar techniques willbe applied to investigate the role of source-sink inter-actions for setting plant species’ range limits in otherstudy systems, such as alpine snowmelt gradients.Research also involves two summer students, JimmyTran and Jen Petzold in a collaborative project withFakhri Bazzaz on the genetic component of intraspe-cific competition in ragweed populations grown atambient and elevated carbon dioxide levels.

Vegetation Dynamics of Ridgetop Pitch Pine and RedPine Communities

Dave Orwig, Glenn Motzkin, and David Foster areconducting a study of the vegetation and long-termdynamics of ridgetop sites that support uncommonpitch pine or red pine communities. Over this pastyear, we focused our efforts on investigating the summit of Mt. Everett in the town of Mt. Wash-ington, Massachusetts. We are using a combinationof dendroecological, historical, and field studies to

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Garlic mustard (Alliaria petiolata): Exotic invader of temperate deciduous understories.

Tree form near the summit of Mt. Everett, southwesternMassachusetts.

address several questions: 1) What is the historicalfire and disturbance history of the region and howhas this influenced vegetation composition and struc-ture? 2) What is the history of land use and how hasthis influenced vegetation composition and structure?3) What is the age structure of the current stand andhow is this related to disturbance history? 4) How dothe composition, structure, and dynamics of Mt.Everett compare with other sites in the southernTaconics and elsewhere in central New England?

Mt. Everett supports unusual dwarf pitch pinesthat are up to ~ 170 years old, but are typically 1–3m (3–10 feet) tall, and occasionally are completelyprostrate. The site experiences frequent damage fromice storms, which presumably contributes to theunusual growth forms. Interestingly, unlike otherdwarf pitch pine barrens, we found little evidence ofrecent fire on Mt. Everett, and no evidence of seroti-nous (closed) cones. Pitch pine has established con-tinuously since the 1830s. Diameter is a poor predic-tor of age in this forest as trees with diameters of only 10 cm differ in age by more than 100 years. Manypines exhibit multiple stems, prostrate growth forms,and evidence of terminal branch damage. Pitch pinering-width patterns displayed extremely slow growth,with many individuals averaging < 0.40 mm yr.–1,and a few growing only 0.08 to 0.30 mm yr.–1 forperiods of up to fifty years. The unusual structure,extremely slow growth rates, and continuous recruit-ment in the absence of recent fire suggest that weath-er and harsh site conditions continue to maintain thisunique community of disturbance–adapted species.

We will continue to examine additional ridgetop

communities (such as Mt. Tom) to place the results ofthis site into a broader landscape context and tomake comparisons with forests that have been influ-enced by fire and other factors.

Historical and Environmental Controls of VegetationComposition

Jesse Bellemare (M.F.S. student) completed work onhis master’s thesis investigating the influence of his-torical land-use on the Rich Mesic Forest communityin western Massachusetts. Rich Mesic Forests are anunusual forest type characterized by sugar mapledominated canopies and species-rich herb layers.These forests typically occur on moist, east-facingslopes over calcium-rich bedrock, such as marble.Results from sites in the towns of Conway andShelburne in Franklin County indicate that the wide-spread forest clearance and agricultural land use ofthe eighteenth and nineteenth centuries has had sig-nificant long-term impacts on the species compositionof Rich Mesic Forests. Forest stands that have devel-oped on abandoned agricultural land typically havelower herb species richness than forest stands thatwere never cleared, due in part to the limited seeddispersal ability of forest herbs associated with thiscommunity.

Forest Policy

Dave Kittredge was on sabbatical from theUniversity of Massachusetts this year, and worked atthe Harvard Forest investigating examples of privateforest landowner cooperation in other countries withtemperate forests and developed economies. Hespent two weeks in Sweden visiting several coopera-tives, and traveled to Oregon with graduate studentand former REU intern Andrew Finley to presentresults of their study of Massachusetts privatelandowners. Working with several student interns,Dave brought to closure an updated map of protect-ed open space in the north Quabbin region, whichforms the basis of a study of conservation potentialamong neighboring landowners and agencies. He hasalso been involved in the data collection of commer-cial timber harvesting throughout Massachusetts, inan effort to characterize its extent and pattern. Davecollaborated with David Foster and Mary Berlik on aHarvard Forest paper that examines the disparitybetween wood consumption and production in theheavily forested state of Massachusetts (The Illusion

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Mt. Tom, a southern Massachusetts trap rock ridge withwhite pine, red pine and pitch pine.

of Preservation: a global environmental argument forthe local production of natural resources).

Timber Harvesting in Massachusetts

To expand the original North Quabbin timber har-vesting study to a statewide level, John Burk hasbeen mapping and collecting data from Chapter 132(the Forest Cutting Practices Act) plans at the variousDepartment of Environmental Management (DEM)offices. To date, approximately 5,200 records havebeen entered from the years 1984–2001, including allstate records from 1997–2001 and virtually all ofeastern Massachusetts records for the entire seven-teen-year period. Data recorded for each mapincludes landowner information, species harvested,acres reported, wetland crossing, and stand treat-ments.

Of the 5,200 plans, approximately 84% were filedby private landowners and 6% by the MetropolitanDistrict Commission, while the remaining 10% weredivided among local and federal agencies, conser-vation groups, sawmills, and the Department ofEnvironmental Management (DEM). The totalacreage reported as harvested was just over 190,000,an average of 37 acres per plan. White pine was themost heavily harvested species, followed by red oak.Harvests generally occurred at similar densities from

the Berkshires through Worcester County, andtapered off strongly as one moved inside the Route I-495 region toward Boston. The next step will be tocomplete the map by adding the rest of the data fromwestern Massachusetts.

Summer student Katie Nicholson has been work-ing on a preliminary analysis of the data in conjunc-tion with her work on socio-economic impacts of log-ging. Once collated and analyzed, we intend to usethe data gathered from this study to explore the effectof timber harvesting as a form of disturbance inMassachusetts, including impacts in highly populatedareas and heavily forested areas.

Botanical Studies

Barry Tomlinson’s research developments includediscoveries on two broad fronts. Calamus is thelargest genus of palms (350 species) and is importantas the natural source of the cane material rattan.Research collaboration with graduate students RenéeRicher and Russell Spangler, and with Jack Fisher atthe Fairchild Tropical Garden, Miami, shows that

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The view to Mt. Wachusett from the Prospect Hill firetower, illustrating the expanse of forest

in central Massachusetts.

A forest harvest in western Massachusetts.

although these climbing palms can develop stems upto 200 m long, their vascular system is discontinuous.This puzzling observation questions assumptionsabout long-distance water transport in plant stemsand is being extended to other climbing palms. Thishas given the opportunity to resurrect unpublishedanalyses carried out by Martin Zimmermann in the1970s.

Stem form in woody plants has long been knownto foresters to include the ability of mature stems toreorient opportunistically by forming peculiar reac-tion fibers (“tension wood fibers”) that have contrac-tile properties. A broadened basis for understandingthis mechanism is provided by unusual distributionof these fibers in a number of tropical plants, oftenwith direct ecological implications. These includeextra-xylary fibers in the tropical gymnospermGnetum that maintain crown form, concentricallydeveloped fibers in the aerial roots of Ficus that makethe “strangling” habit possible, and eccentric fiberdevelopment that erects the seedlings of mangroveRhizophoraceae. The last may explain the distinctiveviviparous condition that characterizes many plantsof mangrove communities.

Harvard Forest LTER Program

The Harvard Forest is one of twenty-five sites in theLong Term Ecological Research (LTER) programsponsored by the National Science Foundation(NSF). Each site addresses questions of a long-termnature; collectively the sites undertake comparativestudies across ecosystems. Representatives from eachsite and NSF meet twice annually to coordinate net-work-wide activities and to collaborate. The centraltheme of the Harvard Forest LTER is interpretationof the structure, composition, and function of forestecosystems in terms of their history of natural andhuman disturbance and environmental change. Thisresearch is being addressed at the stand, landscape,sub-region (e.g., central Massachusetts) and regional(New England) scale.

The research program involves soil scientists,atmospheric chemists, and ecologists studying physi-ological, population, community and ecosystemprocesses. Investigators represent the Department ofBiology (F. Bazzaz, K. Donohue), Earth andPlanetary Sciences (S. Wofsy, B. Munger), andHarvard Forest (D. Foster, D. Kittredge, G. Motzkin,D. Orwig, A. Ellison) at Harvard University, as wellas the Ecosystems Center—MBL, Woods Hole ( J.

Melillo, K. Nadelhoffer, P. Steudler), the ComplexSystems Research Center at the University of NewHampshire ( J. Aber) and the University ofMassachusetts (M. Mulholland). Emery Boose is theLTER data manager with assistance from JuliePallant. The research is organized to maximize theinteractions among scientists from different disci-plines. Four major scientific approaches include: (1)retrospective studies of historical changes in the envi-ronment and ecosystems; (2) long-term measure-ments of forest structure and function; (3) experi-mental manipulations; and (4) synthesis and model-ing. The LTER science group meets approximatelymonthly. The Harvard Forest Ecology Symposium isheld to present current research with abstracts pub-lished annually. The program for the 2001 sympo-sium is included in this report.

Each year, in addition to results generated byHarvard Forest researchers, we highlight studies byour collaborators in the HF LTER Symposium pro-gram that underscore the value of long-term studies.

National Institute for Global Environmental Change(NIGEC)

Harvard University is the Northeastern RegionalCenter for the NIGEC program sponsored by theDepartment of Energy. NIGEC research seeks toimprove the understanding of mechanisms of globalenvironmental change, to develop experimental andobservational programs that enhance the under-standing of ecosystem and regional scale processescontributing to global change, and to provide educa-tional opportunities in global environmental changeresearch. The Center is administered by the Divisionof Applied Sciences at Harvard and most of the fieldstudies are conducted at the Harvard Forest.Researchers include many of the LTER scientists(Bazzaz, Foster, Melillo, Nadelhoffer, Wofsy) in addi-tion to faculty from the State University of New York(D. Fitzjarrald), Woods Hole Research Center (E.Davidson), University of California (S. Trumbore),U.S. Geological Survey (E. Sundquist) and HarvardForest ( J. Hadley).

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HARVARD FOREST ECOLOGY SYMPOSIUM 2002 — ABSTRACTS AND PRESENTATIONS

Titles of Abstracts and Presentations (*summer undergraduate student)

C. Arabia,* E. Davidson, K. Savage, and P. Steudler. Suppression of Soil Respiration by N Deposition.A. Barker Plotkin and D. Foster. Carbon Storage Over 32 Years in a Temperate Deciduous Forest.A. Barker Plotkin, Kristin Wilson,* and D. Foster. Stand Development After Hurricane Disturbance.D. Barnes, J. Elkins and M. McElroy. Greenhouse and Ozone-Depleting Gases in Rural New England.J. Bellemare. An Assessment of Historical Land-Use Effects on Rich Mesic Forests.E. Boose, D. Foster, A. Barker Plotkin, and B. Hall. Hurricane Impacts Across the Yucatan.W. Borken, E. Davidson, K. Savage, P. Steudler, and E. Sundquist. Drought and Snow Effects on Soil Methane

Oxidation.R. Cobb and D. Orwig. Studies of Hemlock Woolly Adelgid Infestation on Decomposition.D. Fitzjarrald, R.. Staebler, R. Saka,i and M. Czikowsky. Forest-Atmosphere Exchange Processes.D. Francis, E. Doughty, and E. Drew. Distribution of Midge (Chironomidae) Remains in New England Lakes. J. Hadley. Old Growth Hemlock: The Annual Pattern of Carbon Exchange.B. Hall, G. Motzkin, D. Foster, M. Syfert, and J. Burk. Historical Land Cover and Land Use in Massachusetts.L. Hutyra, C. Barford, E. Pyle, D. Bryant, B. Curry,* J. Silvis,* and R. Wofsy. Harvesting and Carbon Stocks.S. Kaufman and F. Bazzaz. The Effect of Nitrogen Deposition on Alliaria petiolata and Indigenous Species.D. Kittredge. Forest Preservation to Management: Conservation in a Landscape with Diverse Ownership.D. Kittredge, J. Burk, D. Foster, and G. Motzkin. Timber Harvest as a Form of Disturbance in Massachusetts.M. Kizlinski, D. Orwig, and D. Foster. Vegetation and Ecosystem Response to Hemlock Decline and Logging.K. Lewis. Defense Chemistry in Invasive Plant Species: What Role Does it Play in Invasiveness?H. Lux, F. Bowles, S. Morrisseau, J. Melillo, and P. Steudler. Warming and Ecosystem Carbon Balance.A. Magill and J. Aber. Chronic Nitrogen Additions to Two Forest Stands.P. Micks and K. Nadelhoffer. Soil Respiration and Response to Soil Temperature.Q. Min. Impacts of Aerosols and Clouds on CO2 Uptake over Harvard Forest.Q. Min and B. Lin. Microwave Land Surface Emissivities over Harvard Forest.S. Morrisseau, H. Lux, J. Melillo, P. Steudler, and F. Bowles. Eleven Years of Soil Warming on Prospect Hill.G. Motzkin, D. Foster, B. Hall, and D. MacDonald. Regional-Historical Perspectives into Conservation of

Uncommon Plant Communities: a Study of the Northeastern Coastal Region.J. Munger, C. Horii, A. Bright, J. Budney, et al. Reactive Nitrogen at Harvard Forest: Update for 2001.J. O’Keefe. Regeneration Following Clearcutting of Red Pine Overstory — Year 12.J. O’Keefe and T. Jones. Woody Species Phenology, Prospect Hill Tract, Harvard Forest — 2001.D. Orwig. Forest Dynamics with Chronic Hemlock Woolly Adelgid Infestations in Southern New England.D. Orwig, G. Motzkin, and D. Foster. Ridgetop Pitch Pine and Red Pine in Southern New England.D. Orwig, R. Cobb, M. Kizlinski, and D. Foster. Ecosystem Consequences of Hemlock Woolly Adelgid.T. Parshall, D. Foster, S. Barry Musielewicz, E. Faison, D. MacDonald, and E. Doughty. Fire History in New England

from Charcoal and Pollen in Lake Sediments.L. Sack and N. Holbrook. The “Hydrology” of Temperate Deciduous Leaves.K. Savage, W. Borken, and E. Davidson. Drought Effects on Soil Respiration in a Temperate Forest.K. Stinson and F. Bazzaz. Ecology, Evolution and Physiology of the Invasive Plant Species, Alliaria petiolata.M. Tingley,* D. Orwig, G. Motzkin, R. Field, and D. Foster. Avian Response to Hemlock Decline.S. Trumbore, W. Borken, E. Davidson, and K. Savage. Respired Soil Carbon During Severe Summer Drought.A. Uraguchi. Relationship Between Pattern of Growth/Allocation and Life History of Deciduous Trees.S. Urbanski, C. Barford, J. Munger, and S. Wofsy. Factors Controlling C Exchange: Hourly to Annual Scales.

19

20

2001 LTER AND NIGEC

Ruth Reck. Xihui Lee. Paul Steudler.

Charlie Cogbill, Neal Scott, Neil Pederson. Kim Smith, Dave Orwig, Glenn Motzkin.

Dave Hollinger. Mike McElroy. Kathleen Donohue.

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ANNUAL SYMPOSIUM

Joe Elkins. Dave Fitzjarrald. Julian Hadley.

Neil Pederson.Steve Wofsy.

Bill Munger.Eddy Flux Tower.Lucy Hutyra.

BULLARD FELLOWS

Alison Berry (University of California at Davis)focused her investigations on the effects of intensiveurbanization on natural ecosystem functioning, fromboth scientific and policy perspectives. Her mainemphasis was on soil ecology in relation to urbaniza-tion. In collaboration with the paleoecology group atHarvard Forest, she initiated studies to characterizechanges in soil microbial communities accompanyingland-use changes. She participated in a roundtablesponsored by the Institute for Cultural LandscapeStudies at the Arnold Arboretum on science, policy,and management for urban landscapes.

Betsy Colburn initiated a study on intermittentheadwaters as habitat for stream salamanders andaquatic invertebrates and began developing aresearch program in collaboration with David Orwigand others on potential effects of hemlock woollyadelgid on stream ecology. She continued her vernalpool research by processing field samples from priorcollections, identifying specimens, and verifying iden-tifications at the Harvard University Museum ofComparative Zoology and through consultation withspecialists. Betsy wrote several research papers forpublication, revised a book manuscript, and gavefour scientific presentations. She served as consultantto the Mass. Natural Heritage Endangered SpeciesProfram (NHESP) and The Nature Conservancy(TNC) on issues associated with conservation ofaquatic biodiversity, U.S. EPA on wetlands bioassess-ment, and MA DEM on vernal pool conservation

and management. Betsy served on one graduate stu-dent committee, provided informal mentoring tothree other graduate students, served as judge of stu-dent presentations at North American BenthologicalSociety’s (NABS) annual meeting, and mentored twoREU summer students at Harvard Forest

Aaron Ellison (Mt. Holyoke College) spent themajority of his sabbatical year writing. With col-league Nick Gotelli (University of Vermont), he com-pleted seven papers on the evolutionary ecology ofpitcher plants and the invertebrates that live in asso-ciation with these otherwise carnivorous plants.Notable among these was a paper in Proceedings ofthe National Academy of Sciences on the effects ofnitrogen deposition on pitcher plants, in which theyproposed a simple biological indicator for nitrogendeposition and saturation rates in bogs. He alsobegan work on a biostatistics textbook to be pub-lished by Sinauer Associates.

Colin Orians (Tufts University) initiated a projectexamining how xylem vascular architecture con-strains the response of forest trees to environmentalheterogeneity. In general, trees are thought to be high-ly sectorial — composed of multiple autonomous sec-tors. As a consequence, nutrients captured by one sec-tor are unlikely to travel to other sectors. However,we have found differences among tree species in sec-toriality. Preliminary evidence shows that Betula is notsectorial (resources travel easily throughout theplant), Fraxinus and Quercus are highly sectorial, andAcer shows moderate sectoriality. In addition to quan-tifying patterns of sectoriality, Colin is examining theecological consequences of sectoriality to nutrientcapture and transport. The availability of soilresources is often patchy and Colin is testing thehypothesis that less sectorial species are better able tomove nutrients from resource-rich patches to the mostrapidly growing leaves and branches.

Dr. Akile Pite (Albania) evaluated managementpolicies for protected areas in Albania. To examineU.S. policies, he had contact with the AdirondackPark Agency, the Environmental ConservationCommission in New York State, The Trustees ofReservations, Quabbin Reservoir’s MassachusettsDistrict Commission (MDC) staff, and Friends ofQuabbin. He prepared one manuscript “For BetterManagement of the Protected Areas in Albania.”

Kimberly Smith (University of Arkansas)worked on two edited volumes, Conservation Prioritiesfor Birds at Risk in Latin America and Current Knowledgeand Future Needs for Research on Migratory Birds of

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Betsy Colburn.

Southeastern United States. He also began work on athird volume, The Arkansas Breeding Bird Atlas. Hecompleted manuscripts on habitat mapping for birdconservation in North America, zoogeography ofmammals of the Korean Peninsula, genetic analysis ofblack bears in the southeastern United States, homerange and habitat selection in southern flying squir-rels, human population growth in counties surround-ing federal lands, and potential impacts of the currentred oak borer infestation in the Ozarks on bird com-munity structure. A proposal was submitted to NSFon the last topic in December and resubmitted in July.With Glenn Motzkin, he started a research project onthe history of periodical cicada emergences inMassachusetts and the demise of one brood due toland-use changes in the early to mid-1800s. He pre-sented seminars at the Harvard Forest, BostonUniversity, and Providence College and wrote fourshort historical pieces for the ornithological journal,The Auk, for which he currently is editor-in-chief.

Bernhard Stadler (University of Bayreuth,Germany) initiated an experiment to study the effectsof trophic interactions between hemlock woolly adel-gids and epiphytic bacteria on the spatial and tem-poral variability in flows of energy and matterthrough the canopy of hemlock. At each of three sitesshowing different degrees of infestation (HarvardForest=uninfested, Devils Hopyard=medium infest-ed, Seldom Neck=heavy infested) throughfall solu-tions are collected underneath four trees, each about

5 meters tall. To investigate the within tree variabili-ty in infestation rates, litter production, micro-organism abundance/diversity, and throughfall chem-istry three throughfall samplers were placed beneatheach tree; one close to the trunk, the second close tothe periphery of the canopy, and the third inbetween. The study will be linked to litter decompo-sition experiments by manipulating the quantity andcomposition of wet and dry (e.g., wax) input. Thegoal is to understand the effects of an introduced pestspecies on transforming small-scale ecosystemprocesses and landscapes.

Navjot Sodhi (National University of Singapore)wrote two manuscripts and a review of avian extinc-tions in tropical forests. As tropical forests are beinglost at an alarming rate, his review shows that specieswith large or heavier bodies and those foraging oninsects, fruits, or both are particularly extinctionprone.

Bullard Fellows for 2002–2003 include PhilipBurton (Symbiois Research and Restoration,Canada), Peter Franks (James C. Cook University,Australia), Alice Ingerson (Arnold Arboretum,Harvard University), Matthew Kelty (University ofMassachusetts), David Lindenmayer (AustralianNational University), Francis Putz (University ofFlorida) and Bernhard Stadler (University ofBayreuth, Germany)

EDUCATIONAL ACTIVITIES

David Foster, John O’Keefe, Dave Orwig, GlennMotzkin and other staff members led the HarvardForest Freshman Seminar of eight students in the

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Akile Pite.

Bernhard Stadler.

spring. Barry Tomlinson was facilitator in a course atthe National Tropical Botanical Garden, Kalaheo,Kauai, Hawaii in July 2001, funded by the KenanFoundation. The course objective was to enhance the teaching of botany in introductory biology courses, with an emphasis on tropical plants and wasattended by instructors from ten colleges and universities in North America.

Summer Research Program

The Harvard Forest Summer Student Research pro-gram, coordinated by Edythe Ellin and assisted byLaura Schreeg, attracted a diverse group of studentsto receive training in scientific investigations, andexperience in long-term ecological research. Studentswork closely with researchers and many conducttheir own independent studies. The programincludes weekly seminars with resident and visitingscientists, discussions on career issues in science, andfield exercises on soils, land-use history, and plantidentification. An annual field trip is made to theInstitute of Ecosystem Studies (Millbrook, N.Y.) toparticipate in a Forum on Careers in Ecology.Students present major results of their work at the

Annual Summer Student Research Symposium.

Summer Students 2002

Brent Berger Oklahoma City Univ.Amanda Brow Eastern Kentucky Univ.Brandon Burke University of FloridaMatthew Bur Harvard UniversityElisabeth Dietrich Yale UniversityJacqueline Guzman St. Edwards UniversityJeffrey Law Edinboro UniversityKate Musgrove Ulster County Community

CollegeNicole Nowinski Carleton CollegeSarah Pears Dickinson CollegeJennifer Petzold Trinity CollegeNicholas Povak Virginia Polytechnic Inst. &

State Univ.Gregory Santoni Harvard UniversityJoseph Seggio Skidmore CollegeSigrid Smith Tufts UniversityChristopher Streeter Harvard UniversityJimmy Hung At Tran Pennsylvania State Univ.Naomi Wender Harvard UniversityEugenia Yazzie Northern Arizona Univ.

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Harvard Forest Freshman Seminar.

25

HARVARD FOREST SUMMER STUDENT PROGRAM

Maggie Dietrich.

Jackie Guzman.

Kate Musgrove.

Laura Schreeg, Proctor. Kristina Stinson and Jen Petzold.

Chris Streeter and

Maggie Dietrich.

ACTIVITIES OF THE FISHER MUSEUM

The Fisher Museum plays an important role in theeducational mission of the Harvard Forest by provid-ing a public outlet for research in forest biology, con-servation and management. The Museum also pro-vides a unique setting for conferences and workshopssponsored by the Forest and outside organizations.Dr. John O’Keefe has primary responsibility for thedevelopment of activities and coordination of the useof the Museum.

In late August 2001 the long-awaited exhibitbringing the diorama story up to the present wasinstalled opposite the 1930 land-use diorama. Thisexhibit, consisting of a large, back-lit photograph ofthe modern forest landscape and an adjacent panelillustrating and discussing the many changes in ourvegetation and wildlife that have resulted from theland-use history depicted in the dioramas, was for-mally unveiled at our Annual Meeting in September.

In November the Museum volunteers celebratedthe completion of another successful weekend schedule at our eleventh Volunteer RecognitionDinner, at which Mary Ann Walker received specialthanks for her continuing, enthusiastic work as vol-unteer coordinator. A familiar group, including Billand Marianna Berry, Hector Cameron, WaltDavidson, Bob Lane, and Martha Siccardi, receivedspecial recognition for being the most active volun-teers during the season. In December the group wassaddened by the death of Dr. Richard Riley, one ofour original volunteers and a long-time friend ofHarvard Forest.

During the year the Museum provided programsfor nineteen elementary and secondary schools, thirty-three college and university classes, and twen-ty-two community and professional groups. InDecember the Forest hosted a joint workshop withthe Massachusetts Executive Office of Environ-mental Affairs and the Massachusetts Land TrustCoalition to strategize protection of the Massa-chusetts landscape, and in June the Museum andForest hosted the Massachusetts Department ofEnvironmental Management’s training for summerinterpreters.

Meetings, Conferences, Seminars

The Thirteenth Annual Harvard Forest Long-Term Ecological Research Symposium was held inthe Museum on March 28 followed by the National

Institute for Global Environmental Change meetingon March 29. Other meetings at Harvard Forestincluded meetings of the Massachusetts ExecutiveOffice of Environmental Affairs, MassachusettsExtension Service Coverts Project, MassachusettsDepartment of Environmental Management LoggingWorkshop, Massachusetts Land Trust Coalition,Massachusetts Audubon Society Staff Workshop,Massachusetts Forestry Association, New EnglandForestry Foundation, Mount Wachusett CommunityCollege, National Canon Envirothon PlanningGroup, North Quabbin Regional Landscape Partner-ship, Vegetation Control Service, and the Society ofAmerican Foresters Yankee Division. The Forest alsohosted a faculty retreat for the Harvard UniversityDepartment of Organismic and EvolutionaryBiology.

Speakers in the Harvard Forest Seminar series included:

John F. Ahern U. Mass., AmherstMarco Albani Università di Parma, ItalyAnna Axelsson Swedish University of

Agricultural SciencesJesse Bellemare Harvard ForestAlison Berry University California at

DavisAaron Ellison Mount Holyoke CollegeElizabeth Farnsworth New England Wild Flower

Society Richard Forman Harvard UniversityNick Gotelli University of Vermont

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The boardwalk through Black Gum Swamp provides museum visitors and classes delightful access to

an interesting forested wetland.

Matt Kizlinski Harvard UniversityMatthew Landis Middlebury CollegeJohn Lichter Bowdoin CollegeAndrea Lloyd Middlebury CollegePaul Moorcroft Harvard UniversityDiana Muir Newton, MassachusettsRobert O’Connor MA Executive Office of

Environmental AffairsColin Orians Tufts UniversityLisa Park University of AkronDylan Parry SUNY, SyracusePeter Paton University of Rhode IslandAkile Pite Bullard Fellow, AlbaniaKaren Searcy U. Mass., AmherstKimberly Smith University of ArkansasNavjot Sodhi National Univ. of SingaporeJay Stager Paul Smiths College

FOREST MANAGEMENT AND MAINTENANCE

The most important mission of the Woods Crew is toprovide research support. This year, major projectsincluded purchasing a Scanlift mobile bucket lift vehi-cle capable of reaching seventy-nine feet up into treecanopies with funding from NSF, HU OEB and HFendowment funds; building an equipment shed forJulian Hadley’s new tower; and completely renovat-ing the research scaffold towers, some of which wereover ten years old. Another highlight of the year wasthe major renovation to the facilities portion of thebasement including creation of a spacious newresearch equipment room and a spacious woodwork-ing area with new lighting, an excellent air qualitysystem, and new machinery. This project required

the removal of decades of accumulated old fixturesand research equipment from the basement as well assignificant asbestos abatement.

Another important project focused on the frontsof Shaler Hall and the Fisher Museum. This project

included: expanding and paving the front parkingarea around Shaler Hall; laying out new cobblestonewalkways to the entrances, lighting improvements,and new landscaping including trees, shrub andplants. To maintain these improvements, we havehired a part-time gardener.

Other significant projects included refurbishingthree apartments in the Community House; improve-ments to the cow pasture including building a newshelter for visiting cows and installing a woodenfence around the meteorological station to protect itfrom inquiring cows.

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Adrian Fabos, New Facilities Manager.

Lucas Griffith.

Woody Cole.

COMPUTERS

Major improvements to our computer facilities con-tinued this year with funding from NSF and HarvardForest. Four new Windows 2000 servers wereinstalled, including a Web server, a tape backup serv-er, and two general-purpose servers attached to a360gb RAID disk array. The new servers will pro-vide a home for the Harvard Forest Web page andassociated databases, centralized backup for individ-ual computers, and a means to provide applicationsoftware to users over the network. At the same time,more than fifty desktop computers were convertedfrom Windows NT to Windows 2000, which offersimproved networking and security as well as power-saving features. A new Intranet Web page was created for use by staff, students, and visiting scien-tists. Two general-use computers were installed in theFisher House. Other improvements included a newvoice mail system and conversion by the Universityof our networking equipment from hubs to switches.New purchases included three laptop computers, anew computer projector for the Seminar Room, anda second color laser printer for the Computer Lab.Plans for the coming year include installing a localWindows 2000 domain, upgrading our Web page fordynamic access to databases, and conversion of ourscientific metadata to the new EML (EcologicalMetadata Language) standard.

ACTIVITIES OF THE HARVARD FOREST STAFF

Audrey Barker Plotkin participated in the course,“Forest Resources Management” at the University ofMassachusetts and a workshop on upland forest useby vernal pool species. Audrey also updated a map of

active research at the Forest and began an inventoryof Harvard Forest’s plantations. Jesse Bellemare pre-sented a poster detailing the results of his research atthe ESA.

Emery Boose was elected to serve on the LTER Information Managers Executive Committee(IMEXEC). He attended the LTER InformationMangers and ESA meetings in Madison, Wisconsin,and an IMEXEC meeting in San Diego, as well asNSF-funded workshops on scientific metadata(Phoenix) and wireless networking (Sevilleta). JohnBurk was named a regular contributor to Natural NewEngland magazine, continued to serve as thePetersham town coordinator for Massachusetts Bio-diversity Days, and assisted the Petersham HistoricalSociety with preparations for the towns upcoming250th anniversary. Julian Hadley presented researchresults at an International Union of Forest ResearchOrganizations (IUFRO) meeting, at ESA, and at anAmerican Geophysical Union (AGU) meeting.

David Foster spent three days in Madison,Wisconsin, where he gave the keynote seminar to theEcology Group’s annual graduate student sympo-sium, led discussions with the environmental history,restoration ecology, and The Intergrative GraduateEducation and Research Traineeship (IGERT)groups, and visited with many colleagues andfriends. He joined Bernd Heinrich and Tom Wesselsin a two-day symposium at Bowdoin College pre-senting talks on conservation and landscape history,and gave the keynote seminar at the annual meetingof the Ecotarium in Worcester. In June David waspresented with the Goodwin-Niering Center’sEnvironmental Achievement Award at ConnecticutCollege. He gave seminars and led discussions withstudents at Connecticut College and in Concord,Massachusetts, the latter in conjunction with a talk atthe Concord Land Trust. David and Steward Pickettjoined together to lead seminars and discussions onecological and conservation issues in urban-suburbanareas for the Arnold Arboretum’s Institute ofCultural Landscapes. David represented the Forest atthe LTER Coordinating Committee meeting inAlbuquerque, participated in a (successful) writingworkshop for a NSF biocomplexity proposal, andcontinued to serve as editor for Ecosystems andNortheastern Naturalist, and on the boards forNIGEC, Highstead Arboretum and ConservationResearch Foundation. In July he traveled with hisfamily to the Galapagos where he was a lecturer on aHarvard Alumni Association trip.

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Emery Boose.

Dana MacDonald led six field trips for theUniversity of Massachusetts Department ofLandscape Architecture and Regional Planning, onecommunity outreach woody plant identification tripat the Harvard Forest, and a field trip at the 2001Pine Barrens Research Forum at BrookhavenNational Laboratories, Brookhaven, N.Y. He co-led afield trip for the State of Massachusetts BiodiversityDays in the town of Pelham and worked with AudreyBarker Plotkin to organize the Harvard Forest semi-nars and lab group meetings.

Glenn Motzkin presented seminars at Bridge-water State College, Pelham Historical Society, NewEngland Botanical Club, and The Nature Conser-vancy’s Hoft Farm on Martha’s Vineyard, where hespoke to summer research assistants working for theMarine Biological Laboratory and The NatureConservancy. Glenn attended conferences on Pre-scribed Fire in Massachusetts, Rare Lepidoptera inNortheastern Pine Barrens, and the Ecology andManagement of Early Successional Habitats in theNortheast. Glenn also led field trips at Mt. Tekoa andHarvard Forest for the Freshman Seminar class, aswell as a field trip to Montague Plain for the NewEngland Botanical Club. Glenn served on the thesiscommittees of Els Malfait and Frederik Debaeke, twoundergraduates from Katholieke Universiteit Leuvenin Belgium who spent two months at Harvard Forest.He continues to serve as an Ecology Advisor for TheTrustees of Reservations, an associate member of the

Massachusetts Natural Heritage and EndangeredSpecies Program Advisory Committee, and a member of the Cooper Award Committee of theEcological Society of America.

John O’Keefe gave talks on Harvard Forestresearch and the history of northeastern forests at theUniversity of Massachusetts in Amherst and theEastern Native Tree Society meeting at MohawkTrail State Forest, and led workshops on forest inter-pretation at the Massachusetts Department ofEnvironmental Management interpreter’s training.John again judged the Mahar High School ScienceFair in March, and serves on the boards of the MountGrace Land Conservation Trust, where he is cur-rently vice president, Massachusetts Forestry Associ-ation and Millers River Environmental Center, andon the executive committee of the North QuabbinRegional Landscape Partnership. He also continuesto serve on the Quabbin Science and TechnicalAdvisory Committee and Secretary Durand’s Advi-sory Group on Environmental Education. In MayJohn and his wife, Lynne Stopen, traveled to Chinato meet and bring home their new daughter, Sara.

Dave Orwig attended and presented two talks atthe Hemlock Woolly Adelgid Symposium in EastBrunswick, N.J., and presented a seminar at theUniversity of Massachusetts, where he became anadjunct faculty member.

Julie Pallant attended New England RegionalComputing Program (NERCOMP) 20002 AnnualConference sponsored by EDUCAUSE, “NewDirections in Collaboration: Support, Strategy, Con-tent, Emerging Technologies, and Infrastructure.”She also completed “Web Development and Funda-mentals,” a graduate course at the Harvard ExtensionSchool. Kristina Stinson presented a paper at ESA.

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A trip to the Big Reed virgin forest in northern Maine byDave Orwig, Glenn Motzkin, and David Foster was

guided by Shawn Fraver and other graduate students from the University of Maine.

Glenn Motzkin and Tom Rawinski.

John Aber University of New HampshireMark Ashton Yale UniversityWilliam Bain Harvard UniversityPeter Bakwin Harvard UniversityCarol Barford University of WisconsinDiana Barnes Harvard UniversityFakhri Bazzaz Harvard UniversityK. Boering Harvard UniversityWerner Borken Woods Hole ResearchRichard Bowden Allegheny College Frank Bowles Ecosystems Center — MBLAlfram Bright Harvard UniversityDavid Bryant Harvard UniversityJohn Budney Harvard UniversityLiz Burroughs Ecosystems Center — MBLSean Burrows University of WisconsinMichael Canfield Harvard UniversityV. Y. Chow Harvard UniversityAlan Coleman Harvard UniversityWilliam Currie Virginia Polytechnic Univ.Eric Davidson Woods Hole Research CenterBruce Daube Harvard UniversityFrederik Debaeke Kuleuven, BelgiumKathleen Donohue Harvard UniversityJim Ehleringer University of UtahBob Evans U.S.D.A. Forest ServiceRebecca Field University of MassachusettsDavid Fizjarrald SUNY, AlbanyAndrew Finley University of MassachusettsRichard Forman Harvard UniversitySteven Frolking University of New HampshireJulia Gaudinski UCLA, IrvineElaine Gottleib Harvard UniversityGonzalo Giribet Harvard UniversityMark Helmlinger NASA Jet Propulsion

LaboratoryJoseph Hendricks University of New HampshireMichelle Holbrook Harvard UniversityDavid Hollinger U.S.D.A. Forest ServiceCassandra V. Horii Harvard UniversityLucy Hutyra Harvard UniversityDaniel Jacob Harvard UniversityChristine Jones Harvard UniversitySylvan Kaufman Harvard UnversityOtto Klemm Univ. of New HampshireJuri Knjazikhin Boston University

Takashi Kohyama Hokkaido UniversityChun-Ta Lai University of UtahCathy Langtimm U.S.G.S.; Holy Cross CollegeBarry Lefer Univ. of New HampshireKristin Lewis Harvard UniversityHeidi Lux Ecosystems Center — MBLAlison Magill Univ. of New HampshireLynn Margulis University of MassachusettsMary Martin Univ. of New HampshirePeter Melcher Harvard UniversityJerry Melillo Ecosystems Center — MBLPatricia Micks Ecosystems Center — MBLJacqueline Mohan Duke UniversityJennie Moody University of VirginiaJeff Morisette NASASarah Morisseau Ecosystems Center — MBLMitch Mulholland University of MassachusettsRanga Myneni Boston UniversityJ. William Munger Harvard UniversityKnute Nadelhoffer Ecosystems Center — MBLTommi Nyman Harvard UniversityJeffrey Parker Smithsonian Environ. Res. Jeri Lynn Parrent Duke UniversityJennifer Pontius U.S. Forest ServiceRonald Prinn M.I.T.Elizabeth Pyle Harvard UniversityMichael Rogers GA Institute of TechnologyLawren Sack Harvard UniversityKathleen Savage Woods Hole Research CenterDavid Schaub University of ArizonaTim Sipe Franklin & Marshall CollegeDavid Sonnenfroh Physical Sciences Inc.Rachel Spicer Harvard UniversityRolf Staebler SUNY, AlbanyPaul Steudler Ecosystems Center — MBLBritt Stephens Harvard UniversityKristina Stinson Harvard UniversityEric Sundquist U.S. Geological SurveyRobert Talbot Univ. of New HampshireMatt Thompson Harvard UniversitySusan Trumbore University of CaliforniaMel Tyree University of VermontAya Uraguchi Hokkaido UniversityShawn Urbanski Harvard UniversityKris Verheyen Kuleuven, BelgiumMaciej Zwienecki Harvard University

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VISITING RESEARCH SCIENTISTS AT THE HARVARD FOREST 2001–2002

A large number of Harvard University and outside scientists use Harvard Forest facilities and research sites. Many of these scientists are involved in the Harvard Forest LTER or NIGEC programs.

PUBLICATIONS

Barford,C. C., S. C. Wofsy, M. L.Goulden, J. W.Munger, E. H. Pyle, S. P. Urbanski, L. Hutyra, S.R. Saleska, D. Fitzjarrald and K. Moore. 2001.Factors controlling long- and short-term seques-tration of atmospheric CO2 in a mid-latitude forest. Science 294: 1688–1691.

Bauer, G. A., G. M. Berntson, and F. A. Bazzaz.2001. The effect of elevated CO2 and increasedN availability on regenerating temperate forestcommunities: biochemical versus stomatal limita-tion of photosynthesis. New Phytologist 152:249–266.

Bazzaz, F. A. and K. A. Stinson. 2001. Genetic versusenvironmental control of ecophysiologicalprocesses: some challenges for predicting com-munity responses to global change. Pp. 283–295In M. C. Pres, J. D. Scholes, and M. G. Barker,eds., Physiological Plant Ecology. British EcologicalSociety, U.K.

Bellemare, Jesse. 2002. Environmental and historicalcontrols on the distribution and variation of richmesic forests in western Massachusetts. M.F.S.Thesis, Harvard University.

Berlik, M. M., D. B. Kittredge, and D. R. Foster.2002. The Illusion of Preservation: A GlobalEnvironmental Argument for the Local Production ofNatural Resources. Harvard Forest Paper No. 26,Harvard University, Petersham, Massachusetts.

Boose, E. R., D. R. Foster, A. Barker Plotkin and B.Hall. 2002. Geographical and historical variationin hurricanes across the Yucatan Peninsula. In A.Gómez-Pompa, M. F. Allen, S. Fedick, and J. J.Jiménez-Osornio, eds., Lowland Maya Area: ThreeMillennia at the Human-wildland Interface. New York:Haworth Press.

Cavender-Bares, J.,M. Potts, E. Zacharias and F. A.Bazzaz. 2001. Consequences of CO2 and lightinteractions for leaf phenology, growth andsenescence in Quercus rubra. Global Change Biology6: 877–887.

Currie, W. S., K. J. Nadelhoffer, and B. Colman.2002. Long-term movement of 15N tracers intofine woody debris under chronically elevated Ninputs. Plant and Soil 238: 313–323.

Dail, D. B., E. A. Davidson and J. Chorover. 2001.Rapid abiotic transformation of nitrate in an acidforest soil. Biogeochemistry 54; 131–146.

Donohue, K. 2002. Germination timing influencesnatural selection on life-history characters in

Arabidopsis thaliana. Ecology 83: 1006–1016.Fisher, J. B. and P. B. Tomlinson. 2002. Tension

wood fibers are related to gravitrophic movementof red mangrove (Rhizophora mangle) seedlings.Journal of Plant Research (Japan) 115: 39–45.

Fitzjarrald, D. R., O. C. Acevedo, and K. E. Moore.2001. Climatic consequences of leaf presence inthe eastern United States. Journal of Climate 14:598–614.

Johnson, T. C., E. T. Brown, J. McManus, S. Barry,P. Barker, and F. Gasse. 2002. A high-resolutionpaleocimate record spanning the past 25,000years in southern East Africa. Science 296:113–132.

Kizlinski, M. 2002. Vegetation and ecosystemresponse to eastern hemlock decline and logging:direct and indirect consequences of the hemlockwoolly adelgid. M.F.S. Thesis, Harvard Uni-versity.

Motzkin, G., S. C. Ciccarello, and D. R. Foster. Frostpockets on a level sand plain: does variation inmicroclimate help maintain persistent vegetationpatterns? Journal of the Torrey Botanical Society 129:154–163.

Motzkin, G., D. A. Orwig, and D. R. Foster. Historyand dynamics of a ridgetop pitch pine community, MountEverett, Massachusetts. Harvard Forest Paper No.25, Harvard University, Petersham, Mass-achusetts.

Savage, K. E. and E. A. Davidson. 2001. Interannualvariation of soil respiration in two New Englandforests. Global Biogeochemical Cycles 15: 337–350.

Sipe, T. W. and F. A. Bazzaz. 2001. Shoot damageeffects on regeneration of maples (Acer) across anunderstory-gap microenvironmental gradient.Journal of Ecology 89: 761–773.

Tomlinson, P. B., and R. Spangler. 2002.Developmental features of the discontinuousstem vascular system in the rattan palm Calamus(Arecaceae-Calamoideae-Calamineae). AmericanJournal of Botany 89: 1128–1141.

Turner, B. L. II, S. C. Villar, D. Foster, J. Geoghegan,E. Keys, P. Klepeis, D. Lawrence, P. M.Mendoza, S. Manson, Y. Ogneva-Himmelberger,A. B. Plotkin, D. P. Salicrup, R. R. Chowdhury,B. Savitsky, L. Schneider, B. Schmook, and C.Vance. 2001. Deforestation in the southernYucatan peninsular region: an integrativeapproach. Forest Ecology and Management 154:353–370.

Williams, C. G., Y. Zhou, and S. E. Hall. 2001. A

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chromosomal region promoting outcrossing in aconifer. Genetics 159: 1283–1289.

Wilson, K. 2002. Thirty-two years of populationchange in a hardwood stand at Harvard Forest,Petersham, Massachusetts. Senior Thesis, Middle-bury College.

Wofsy, S. C. 2001. Where has all the carbon gone?Science 292: 2261–2263.

ACKNOWLEDGMENT OF SUPPORT

Activities described in this Annual Report are sup-ported by funds provided by the following sources, towhich we are indebted:

Friends of the Harvard ForestCommonwealth of Massachusetts

Department of Environmental Management

Division of Fisheries and WildlifeMassachusetts Biodiversity InitiativeEcological Restoration ProgramMassachusetts Environmental TrustMassachusetts Natural Heritage and

Endangered Species ProgramAndrew W. Mellon FoundationDepartment of Energy, National Institute

for Global Environmental ChangeNational Science Foundation

Biological Field Stations and Marine Laboratories

Digital Government ProgramEcosystem StudiesInternational ProgramsLong-term StudiesEcology ProgramResearch Experience for

UndergraduatesThe Family of Richard Thornton FisherUSDA Competitive Research Grants,

ForestrySouthern Taconics Research and

Conservation Center

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GIFTS

Donations to the Harvard Forest library from JimBaird were a welcome addition to its journal collec-tion. Two years of The Auk and Journal of FieldOrnithology were added. Bob Willhite donated a copyof his Harvard University Master of Forest Sciencethesis “Structure and Interrelationships of NewEngland Attitudes and Perception Towards SomeAspects of Forest Management.” Also Dave Kittredgegave a copy of “Southern Forest ResourceAssessment Summary Report.”

NEW FUNDING

Betsy Von Holle and David Foster. Disturbance historiesas a predictor of habitat invasibility in a mosaic landscape:Cape Cod National Seashore. National Park Foundation.$150,000

David Foster and Kathleen Donohue. Harvard ForestREU research program in forest ecology. HarvardUniversity Program for Training and Research in Ecology.National Science Foundation. $237,918.

Julian Hadley and David Foster. Harvard Forest carbonexchange modeling work in connection with digital aerialimage analysis by U. Mass., Amherst. National ScienceFoundation. $10,000.

David Foster and Michelle Holbrook. A mobile canopyaccess platform for environmental and physiologicalresearch at Harvard Forest. National Science Foundation.$55,860.

David Foster and Julian Hadley. ITR/AP: The analyticalweb. (With U. Mass.). National Science Foundation.$300,000.

David Foster. Harvard University program for trainingand research in ecology. Andrew W. Mellon Foundation.$500,000.

Elizabeth Colburn and David Foster. Vernal pools in alandscape context: biodiversity and habitat. MassachusettsEnvironmental Trust. $20,000.

Elizabeth Colburn and David Foster. Monitoring and bio-logical assessment of headwater habitat streams. EOEA-Massachusetts Environmental Trust. $19,924.

Julian Hadley and D. R. Foster. Evaluating the effects ofdiverse vegetation types and soil drainage classes on netcarbon exchange of a landscape mosaic with mobile andfixed eddy covariance systems. NIGEC. $153,335.

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David R. Foster Petersham, MassachusettsDirector August 2002