THE HARVARD FOREST 2000–2001Harvard University

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Personnel at the Harvard Forest - - - - - - 2Introduction to the Harvard Forest - - - - - - 3New Staff - - - - - - - - - 4Research Activities - - - - - - - - 4Bullard Fellows - - - - - - - - - 20Educational Activities - - - - - - - - 21

Summer Research Program - - - - - - 22Activities of the Fisher Museum - - - - - - - 23

Meetings, Seminars, Conferences - - - - - - 23Forest Management and Maintenance - - - - - - 24Long-Term Data Management - - - - - - - 24Activities of the Harvard Forest Staff - - - - - - 26Visiting Research Scientists - - - - - - - 28Publications - - - - - - - - - 29Acknowledgment of Support - - - - - - - 31Gifts - - - - - - - - - - 32New Funding - - - - - - - - - 32

ANNUAL REPORT OF THE HARVARD FOREST, 2000–2001

Rebecca Anderson M.F.S. CandidateAudrey Barker Plotkin Research AssistantSylvia Barry Musielewicz Research AssistantGutram Bauer Post-doctoral FellowJesse Bellemare MFS CandidateEmery Boose Information and

Computer ManagerWerner Borken Post-doctoral FellowJeannette Bowlen AccountantNick Brokaw Bullard FellowJohn Burk ArchivistSusan Clayden Research AssistantRichard Cobb Research AssistantWillard Cole Woods CrewKathleen Donohue Bullard FellowElaine Doughty Research AssistantRobert Eberhardt MFS CandidateJohn Edwards Forest ManagerEdythe Ellin AdministratorEd Faison Research AssistantMary Ann Fajvan Bullard FellowSamantha Farrell Laboratory TechnicianBarbara Flye Librarian/SecretaryCharles H. W. Foster AssociateDavid Foster DirectorDonna Francis Research AssociateJanice Fuller Research AssociateKelli Graves Secretarial AssistantJulian Hadley Research AssociateBrian Hall Research AssistantLinda Hampson Secretarial AssistantJon Harrod Post-doctoral FellowDonald Hesselton Part-time Woods

Crew

Malcolm Hughes Bullard FellowSusan Johnson Research AssistantTeresa Jones Assistant Schoolyard

CoordinatorDavid Kittredge Forest Policy AnalystMatt Kizlinski MFS CandidateOscar Lacwasan CustodianErin Largay Research AssistantSarah Laubscher Summer Program

AssistantManuel Lerdau Bullard FellowDana MacDonald Research AssistantLisa Marselle Summer CookLucinda McWeeney Bullard FellowLaurie Miskimins Research AssistantFraser Mitchell Bullard FellowGlenn Motzkin Plant EcologistJohn O’Keefe Museum CoordinatorDavid Orwig Forest EcologistJulie Pallant Assistant Information

and Computer Mgr.Tim Parshall Post-doctoral FellowDorothy Recos-Smith Staff AssistantLauren Sack Post-doctoral FellowJessica Schedlbauer Research AssistantKarn Deo Singh Bullard FellowCharles Spooner Woods CrewFred Swanson Bullard FellowMindy Syfert Research AssistantP. Barry Tomlinson E. C. Jeffrey Professor of

BiologyJohn Wisnewski Woods CrewSteven Wofsy Associate

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PERSONNEL AT THE HARVARD FOREST 2000–2001

INTRODUCTION TO THE HARVARD FOREST

Since its establishment in 1907 the Harvard Forest hasserved as a center for research and education in forestbiology. Through the years researchers have focussedon silviculture and forest management, soils and thedevelopment of forest site concepts, the biology oftemperate and tropical trees, forest ecology, forest eco-nomics, landscape history, conservation biology, andecosystem dynamics. Today, this legacy of researchand education continues as faculty, staff, and studentsseek to understand historical and modern changes inthe forests of New England and beyond, resultingfrom human and natural disturbance processes, andto apply this information to the conservation, man-agement, and appreciation of natural ecosystems.This activity is epitomized by the Harvard ForestLong Term Ecological Research (HF LTER) pro-gram, which was established in 1988 through fundingby the National Science Foundation (NSF).

Physically, the Harvard Forest is comprised ofapproximately 3,000 acres of land in the north-centralMassachusetts town of Petersham that include mixedhardwood and conifer forests, ponds, extensivespruce and maple swamps, fields and diverse planta-tions. Additional land holdings include the 25-acrePisgah Forest in southwestern New Hampshire (locat-ed in the 5000-acre Pisgah State Park), a virgin forestof white pine and hemlock that was 300 years oldwhen it blew down in the 1938 Hurricane; the 100-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 Laboratoriesprovide office and laboratory space, computer andgreenhouse facilities, and a lecture room for seminarsand conferences. Nine additional houses provideaccommodations for staff, visiting researchers, andstudents. Extensive records, including long-term datasets, historical information, original field notes, maps,photographic collections, and electronic data aremaintained in the Harvard Forest Archives.

Administratively, the Harvard Forest is a depart-ment of the Faculty of Arts and Sciences of HarvardUniversity. The Harvard Forest administers theGraduate Program in Forestry that awards a Mastersdegree in Forest Science and faculty at the Forestoffer courses through the Department of Organismicand Evolutionary Biology, the Kennedy School of

Government, and the Freshman Seminar Program.Close association at Harvard is also maintained withthe Department of Earth and Planetary Sciences , theSchool of Public Health, and the Graduate School ofDesign, and with the Department of NaturalResource Conservation at the University of Massa-chusetts, the Ecosystems Center of the MarineBiological Laboratory at Woods Hole, and the Com-plex Systems Research Center at the University ofNew Hampshire.

The staff and visiting faculty of approximatelyfifty work collaboratively to achieve the research,educational and, management objectives of theHarvard Forest. A management group comprised ofthe Director, Administrator, Coordinator of theFisher Museum, and Forest Manager meets monthlyto discuss current activities and to plan future pro-grams. Regular meetings with the HF LTER scienceteam, weekly research seminars and lab discussions,and an annual ecology symposium provide for aninfusion of outside perspectives. The four-memberWoods Crew and Forest Manager undertake forestmanagement and physical plant activities. TheCoordinator of the Fisher Museum oversees manyeducational 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, and the AndrewW. Mellon Foundation. Our summer Program forStudent Research is supported by the NationalScience Foundation, the A. W. Mellon Foundation,and the R. T. Fisher Fund.

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NEW STAFF

Teresa Jones who received her M.A. in plant biologyfrom UMass-Amherst is the new Assistant LTERSchoolyard Coordinator, involving K through 12teachers and students in long-term research projects.Jessica Schedlbauer, a graduate of the University ofMaine with an M.S. in forest ecology, is working as aresearch assistant for Julian Hadley on the NIGECproject. Kelli Graves is a new office assistant andreceptionist at the Forest.

RESEARCH ACTIVITIES

An Historical Approach to Understanding andConserving the Coastal Region

A major goal of plant ecology is to determine the fac-tors that control species distributions and communi-ty composition through time and space. Althoughvegetation patterns result in part from species-specif-ic responses to environmental conditions andresource availability, actual species distributions atany point in time differ from potential distributionsas a result of several factors, especially biotic interac-tions and the history of natural and human distur-bance. Disturbance may influence community pat-

terns by (1) directly altering the environment andresource distributions; (2) creating opportunities forthe establishment of new species; or (3) reducing pop-ulations of established species. For instance, our pre-vious work has demonstrated that species with slowrates of dispersal or establishment may be absentfrom a site for decades or centuries, not because thesite is inherently unsuitable, but simply because thespecies were removed by prior disturbance and havenot had sufficient time to recolonize. Consequently amajor challenge for ecological study is to evaluate therelative contribution of current environmental condi-tions and historical factors in determining modernvegetation patterns and dynamics.

Following up on our previous studies in centralMassachusetts, the Connecticut Valley, and Martha’sVineyard, we are conducting a comprehensive inves-tigation into 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 andanimal species and communities and because it ishighly threatened by development. With supportfrom the National Science Foundation, MellonFoundation, Massachusetts Biodiversity Initiative,and The Nature Conservancy’s Ecological Research

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Cape Cod showing major details of the suficial geology and location of pollen analyses (left) and woodland cover in the nineteenth and twentieth centuries (right).

Program, we are investigating the link between thelandscape history and environment of the coastalregion and the modern abundance and distributionof upland plant communities, including grasslands,heathlands, barrens, and woodlands. Although theregion has a long history of settlement and intensiveuse, there has been no prior attempt to evaluate rig-orously the impact of land-use history on species dis-tributions and community assemblages. Such aregional understanding is critical in order to interpretmodern landscape and species patterns, to identifyecological goals for conservation, and to developmanagement approaches for achieving those objec-tives.

Our current work emphasizes integrated paleoe-cological, historical, Geographic InformationSystems (GIS), and field studies throughout thecoastal region. Tim Parshall and David Foster areconducting paleoecological studies to reconstruct thelong-term vegetation and disturbance history of theregion, and Brian Hall is gathering a wide range ofhistorical data, including excellent maps for incorpo-ration into a growing GIS database. This past year

Brian and John Burk researched historical records,including farm journals and diaries, town records,maps, and publications on Nantucket. A summary oftown acts regulating agriculture on the island wascompiled, and copies were made of old documents.Rob Eberhardt and Jon Harrod worked closely withDavid and Glenn Motzkin to evaluate the relation-ship between modern vegetation variation, site fac-tors, and disturbance history. This past year, Rob fin-ished his Master’s thesis on the historical influenceson modern forest vegetation on Cape Cod NationalSeashore. More than half of the National Seashorewas cleared for agriculture in the nineteenth century,and although these agricultural lands have long beenabandoned, Rob documented substantial differencesin vegetation composition between forests on oldfields and those on areas that were never cleared.This summer, Dana MacDonald, Lynda Joudrey,and Tricia Burgoyne are sampling forests on LongIsland and Martha’s Vineyard to provide a broaderunderstanding of factors controlling vegetation varia-tion across the coastal region.

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Species distributions in forests on Cape Cod NationalSeashore are strongly controlled by history, in particularwhether the site was continuously woods or subjected toforest clearance, agricultural plowing, and subsequentreforestation.

Brent Wolfe, Tim Parshall, and Sylvia Barry Musielewicztaking a sediment core.

While the modern and historical studies are pro-viding a detailed picture of ecological changes afterthe arrival of Europeans, the paleoecological researchis uncovering how vegetation, fire, and the environ-ment changed over the past several thousand years.This work, headed by Tim Parshall, also provides acontext for interpreting the relative effects of humanversus natural factors in driving more recent land-scape dynamics. In this study Ed Faison, DanaMacDonald, and Elaine Doughty are looking at fos-sil pollen and charcoal preserved in the sediments ofa series of lakes arrayed so as to sample major varia-tion in geography, soils, and land use across theregion. European settlement on Cape Cod in the1600s shows up clearly in these records as a rise inthe amount of pollen from grasses and herbs, reflect-ing the clearance of forests for pastures and agricul-tural fields.

Before European settlement, fire and vegetationpatterns were closely linked with physiography, soils,and geography. Oaks and other hardwoods weremore abundant on moraines, and pitch pine and firewere more abundant on outwash. Fire was also morecommon on the western Cape, which is broader andmore susceptible to the development of large fires.There is no evidence for Indian clearing or agricul-ture. In contrast, the modern vegetation is lessdiverse, more regionally homogeneous, and exhibitsless variation with the factors important beforeEuropean settlement. These results are importantbecause they highlight the heterogeneous nature of the early landscape and vegetation and illustrate

some of the biotic and physical characteristics thatmay be targets for conservation and restoration, Thisresearch has also developed ways to reconstruct and compare fire histories across New England.Results indicate that before the arrival of Europeansfire was most common on Cape Cod and coastalNew England and was much less important in northern hardwood forests in central Massachusetts.Isolated areas with higher fire existed, including thepitch pine-oak forests of the Connecticut Valley.

In a related coastal study Susan Clayden completed a pollen diagram for Harlock Pond onSeven Gates Farm on the western side of Martha’sVineyard. The site is surrounded by open deciduousforest and was once owned by Nathaniel Shaler, professor and dean at Harvard and an original supporter of the concept of a “Harvard Forest.” The 2,500-year record indicates that oaks, beech, and birch have been the dominant trees, that fire was uncommon, and, like many of our other records,that there was a significant shift in vegetation 1,500years ago. Through the past 300 years of Europeanactivity there has been an increase in blackgum (Nyssasylvatica), heaths (Ericaceae), and sweet pepperbush(Clethra alnifolia), accompanied by a decrease in beechand hemlock. Pollen of Atlantic white cedar was alsomore abundant prior to settlement.

The Little Ice Age in New England

David Foster, Donna Francis, and Janice Fuller arecoordinating a study of changes associated with the

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Multivariate analysis of all pollensamples from seven sites onCape Cod. Before European set-tlement the vegetation exhibited compositional differences where-as, samples from the seventeenthcentury to the present indicate amuch more homogeneous vege-tation as a result of land-useimpacts.

Little Ice Age, a period of cooler and variable climatefrom approximately 1400–1850 A.D., and humanactivity. Understanding the factors driving vegetationchange that occurred before and following the influ-ence of European settlement is essential for interpret-ing current vegetation patterns and developing long-term conservation policies for the modern landscape.Signals of climate change, fire, human and naturaldisturbance, and vegetation change are being investi-gated in a multiproxy study of eight lakes on a north-south transect from northern Vermont to southernConnecticut. Results from this study will be inte-grated with those from the coastal project to providea New England-wide perspective.

Dating, pollen analysis, and chironomid analysishave been completed for six sites. Dörte Köster atLaval University is analyzing sedimentary diatomsand modern diatom assemblages at two sites. NatalieDrake, Susan Clayden, Ed Faison, and DanaMacDonald are doing pollen analysis, and ElaineDoughty and Donna Francis are analyzing chirono-mids. Brent Wolfe at the University of Waterloo willanalyze stable isotopes. Preliminary results of chirono-mid analyses from 6 ponds indicate a temperature shiftof 1.3° C during the time of the Little Ice Age in NewEngland. Two more ponds in Vermont andMassachusetts will be cored to complete the study.

Sylvia Barry Musielewicz coordinated daily activities for the paleoecology lab and participated infieldwork to recover sediment cores for the Little IceAge and Cape Cod projects.

Massachusetts Forest Composition and Distributionthrough Time: Providing the Context for Ecologicaland Biodiversity Assessment

Because Massachusetts has a long cultural history,ecological study, conservation, and long-term plan-ning must incorporate an understanding of the pastland-use and landscape modification. However, todate, most ecological studies and planning activityhave failed to evaluate human history, primarilybecause information at local and regional scales hasbeen unavailable. With support from theMassachusetts Natural Heritage and EndangeredSpecies Program, David Foster, Glenn Motzkin,Mindy Syfert, and Brian Hall are developing GISdata layers that will allow for rapid assessment of thehistory of intensive land use throughout theCommonwealth. Using maps that were generated foreach town in 1830, we are developing maps indicat-ing forest land, open land, meadows, and cultural fea-tures, including roads, mills, meeting houses, etc.Because 1830 was near the period of maximum for-est clearance, the resulting data layers will provide areasonable approximation of the Massachusetts land-scape at the height of agricultural activity. Such infor-mation will be valuable for research, planning, andmanagement, including evaluation and prioritizationof land for conservation, restoration, and long-term

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Ed Faison.

Brian Hall and Mindy Syfert.

ecological management, comparison of primary ver-sus secondary woodlands, and archaeological andcultural resource assessment. Interestingly, the mapsshow that the greatest extent of forestland inMassachusetts in 1830 occurred in southeasternMassachusetts, where large portions of Plymouthand Bristol Counties and inner Cape Cod remainedforested. In contrast with the modern landscape,isolated woodlots in a predominantly agriculturallandscape characterized the uplands of central and western Massachusetts.

Other databases were created from historicalrecords for every town in Massachusetts to evaluatecontrols on vegetation composition during this time.The largest was a detailed summary table of all agri-cultural and woodland data in the state census andUMass/MacConnell land-use records from 1801 tothe present. From the 1885 state census, a separatetable was generated for forest cutting data by species.Other databases were created for the early twentiethcentury county forest surveys, the 1907 stateforester’s report, and historical population.

John Burk worked with David, Glenn, and Brianto gather data on forest composition in southern New

England at the time of European settlement byresearching surveyor records across Massachusetts,Connecticut, and Rhode Island. All but one surveywere proprietor or deed lot layouts, which providedetailed species information. Survey dates rangefrom the 1620s for the coast and Connecticut Valleyto 1830s for interior hill towns. In Connecticut,44,000 trees were tallied covering three-quarters ofthe towns, with gaps in Hartford and Fairfield coun-ties where trees were rarely used in surveys. InRhode Island 85 percent of towns had useful records,resulting in a tally of 2,700 trees and a gap in sixsoutheastern coastal and island towns. InMassachusetts, the total was 58,000 trees from 84percent of the 351 towns. Several towns in the regionhad over 3,000 trees, with a high of 4,500 inKillingworth, Connecticut. These data will provide adetailed understanding of early forest composition insouthern New England as well as changes that result-ed from historical land use and climate change. Inthis project we are collaborating with Charlie Cogbillat Hubbard Brook who has similar data for northernNew England to evaluate forest composition acrossthe entire region.

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Rebecca Anderson, Jesse Bellemare, and Glenn Motzkin.

In these and related studies Brian Hall continuedsupporting our role for GIS activities. He generatedfigures for presentations and publications, analyzedspatial data with a GIS, and worked on systemsdevelopment. As GIS becomes increasingly impor-tant in research, we have added a second digitizingtable to speed conversion of paper maps into digitalformat. This past year we have also purchased ahigh-quality Global Positioning Satellite (GPS)receiver, which allows determinations to within a fewmeters; this will be helpful in locating and relocatingstudy sites as well as mapping Harvard Forest properties.

Forest History and Modern Species Distributions: AComparison of Plant Life History Characteristicsfrom Europe and Eastern North America

Glenn Motzkin and David Foster are collaboratingwith Kris Verheyen, Olivier Honnay, and MartinHermy from the University of Leuven in Belgium ona comparative study of the life history characteristicsof plant species whose current distributions arestrongly linked to historical land-use practices. InEurope and eastern North America, several speciesare characteristic of primary woodlands that werenever cleared for agriculture, but are missing in sec-ondary woodlands that developed on former agricul-tural lands. Although the floras on the two continentsdiffer substantially, our analyses are evaluatingwhether particular plant traits (e.g., number of seeds,dispersal mechanism, etc.) are characteristic ofspecies that are slow to colonize after disturbance.

Hemlock Woolly Adelgid Impacts on New EnglandForests: Changes in Forest Structure, EcosystemFunction, and Bird Communities

This summer Dave Orwig, Richard Cobb, and sum-mer students Katie Theoharides and Spencer Meyerresampled permanent plots on eight sites establishedin 1997 to examine the effect of hemlock decline andmortality on the timing and extent of nitrogen cyclingchanges associated with the introduced hemlockwoolly adelgid (HWA). Soil analyses including pH,temperature, carbon to nitrogen ratios, texture, andtotal soil organic matter were completed and meas-urement of the rates of nitrogen mineralization in asubset of sites is ongoing. After several years of HWAinfestation, tree crowns became damaged, allowinggreater light to reach the forest floor, such that stands

with heavy infestations had lower surface soil mois-ture levels, higher soil temperatures, and higher netnitrogen mineralization and net nitrification ratesthan slightly damaged or uninfested stands. Resinbags buried in the soil at damaged sites also capturedhigher amounts of ammonium and nitrate than inuninfested stands, indicating that nitrogen is becom-ing more available as a result of the microenviron-mental changes associated with HWA damage.Sampling will continue to examine the long-termchanges in nitrogen (N) cycling associated with hem-lock mortality and replacement. Dave and Katie

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Seasonal nitrogen mineralization in healthy hemlockstands, HWA damaged stands, and new and old hemlockharvest sites (upper). Mobile nitrogen in soil solution cap-tured in buried resin bags (lower). Note different scales.Higher nitrification rates are common in disturbed ecosys-tems where nutrient cycles have been altered. A lack ofvegetative uptake in new harvests allows more nitrogrento move deeper into the soil, possibly leaving the system.Despite high nitrification rates, little nitrogen is leaving theupper soil horizons in old harvests.

resampled plots established in 1995 to examine foreststructure changes associated with HWA. Hemlockcontinues to deteriorate in all stands and black birchsaplings in several stands are three to five meters tall.

Last fall Richard Cobb, Dave, and former REUstudent Steve Currie, completed an eighteen-monthstudy of HWA impacts on foliar decomposition.Litter decomposition is a major pathway of nutrientcycling and organic matter accumulation in forestsoils and has a strong influence on forest productivi-ty and soils. Decomposition of hemlock foliage wasmeasured at eight sites with varying degrees of HWAdamage. Stands with substantial canopy damage hadlower rates of decomposition as a result of warm, drysurface soil conditions, a finding that contradictedour hypothesis that increased soil temperature wouldaccelerate decomposition. However, dry soil condi-tions retard the establishment of fungal decomposercommunities in the surface soil. Microbes and fungalcommunities are a large sink of N in healthy forestsand reduced microbial establishment or biomass maybe contributing to N losses in these damaged forests.To expand this investigation we are comparingdecomposition rates at the soil surface versus theorganic-mineral soil interface at twelve sites inConnecticut and Massachusetts. We have also beguna reciprocal litter study comparing hemlock, black

birch, and mixed-litter decomposition. Black birch isthe most prolific species establishing as hemlocksdeteriorate and its foliar chemistry is substantiallydifferent from that of hemlock. Comparing speciesdifferences and potential species interactions willimprove our understanding of how ecosystemprocesses change with hemlock decline.

As a follow-up to last year’s pilot study, BeckyField and returning summer student Morgan Tingleyinvestigated bird species richness and compositionchanges associated with hemlock deterioration fromHWA. Several species including black-throated greenwarblers, hermit thrushes, and blue-headed vireosexhibit significant declines with the loss of hemlock,while tufted titmice, red-eyed vireos, and Americangoldfinches increased as hemlocks were replaced byhardwoods.

Vegetation Dynamics of Ridgetop Pitch Pine and RedPine Communities

As part of a broader investigation of ridgetop com-munities in the Northeast, Dave Orwig, GlennMotzkin, and David Foster initiated a study examin-ing the history, development, and vegetation dynam-ics of Mt. Everett and nearby summits in the south-ern Taconic region of southwest Massachusetts. We

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Jessica Schedlbauer adjusting photosynthesis equipment in the hemlock canopy tower.

are using a combination of dendroecological, histori-cal, and field studies to address several questions: (1)What is the history of fire, natural disturbance, andhuman activity and how has this influenced vegeta-tion composition, structure, and age distribution?And (2) How do the composition, structure, anddynamics of Mt. Everett compare with similar sitesacross central New England?

We have visited several sites and intensively sampled the dwarf pitch pine community on Mt.Everett. This rare forest type, occurring in only a fewlocations in the northeastern U.S., is characterized byexposed bedrock, shallow soil depths of 10 to 18 cm,and abundant stunted pitch pine 0.5 to 2.0 m tall.The uneven-aged stand includes red maple, red oak,and birches. Pitch pine exhibits continuous recruit-ment since the 1830s and ranges from 12 to 170 yearsold. Hardwoods were established in most decadessince 1860 and in higher densities in the 1950s and1960s. Diameter is a poor predictor of age as treeswith diameters of only 10 cm differ in age by morethan 100 years. Many pines exhibit multiple stems,prostrate growth forms, and evidence of terminalbranch damage. There is no historical or visual evidence of fire or human impacts in this forest, buton this exposed summit (795 m a.s.l.), wind, snow,and ice damage are dominant disturbance factors.Pitch pine ring-width patterns displayed extremelyslow growth, with many individuals averaging < 0.40mm yr–1, and a few growing only 0.08 to 0.30 mmyr–1 for periods of up to fifty years. The unusualstructure, extremely slow growth rates, and continu-ous recruitment in the absence of recent fire suggestthat weather and harsh site conditions continue tomaintain this unique community of disturbance-adapted species.

Forest Policy

Dave Kittredge continued exploring patterns of landprotection, forest owner attitudes, and forest man-agement in the North Quabbin landscape of centralMassachusetts. Research on owner attitudes andbehaviors expanded this year with the acquisition ofall private forest management plans for NorthQuabbin properties enrolled in Chapter 61, the forestproperty tax program. When added to the map ofother state lands under management, and the previ-ously acquired data on harvest activity, an improvedview of the extent and type of forest managementwill be developed for this diverse sample region.

Carbon Exchange of Old-growth Hemlock Forests

This fall Julian Hadley initiated long-term measure-ments of carbon exchange in the old-growth hemlockforest in the center of the Prospect Hill tract. Initialresults indicate that these forests stored substantialamounts of carbon in October and early November,but negligible amounts thereafter through mid-March. Carbon storage ceased on November 20 fol-lowing the first daily minimum temperature below–5°C. This result corresponds well with measure-ments of photosynthesis on individual hemlockshoots in previous years, which showed a sharpdecline after nights below –2° C and near-zero pho-tosynthesis after a minimum of –8° C. Such a lowrate of photosynthesis could easily be balanced byrespiration of the soil and trees, leading to no net car-bon storage. Although the evergreen hemlock forestdid not show any net carbon storage in winter, car-bon storage in the fall after leaf drop by deciduoustrees amounted to about 15 percent of average annual carbon storage in nearby deciduous forests.Clearly evergreen trees can derive substantial benefitfrom photosynthesis in mid- to late autumn.

Ecological Impacts of Hurricanes Across the YucatanPeninsula

The ecological impacts of hurricanes across theYucatan Peninsula in Mexico were investigated byEmery Boose, David Foster, Audrey Barker Plotkin,and Brian Hall, using a simple meteorological model(HURRECON) developed at the Harvard Forestand a database of historical hurricane data (HUR-DAT) maintained by the U.S. National HurricaneCenter. A total of 105 hurricanes over the period1851–2000 were reconstructed to produce estimatesof wind damage on the Fujita scale (a common scaleof damage for tornadoes and hurricanes) across theregion. Results showed considerable annual anddecadal variation in hurricanes, while at the half-cen-tury scale there was an increase in hurricane intensi-ty since the mid-nineteenth century. Ninety percentof the hurricanes causing F1 damage or higher (i.e.,more than scattered individual windthrows) occurredin the months of August, September, and October. Astrong spatial gradient in hurricane frequency andintensity extended across the region from northeastto southwest, resulting from the greater number ofhurricanes to the north, the east to west movement ofmost hurricanes across the area, and the tendency for

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most hurricanes to weaken significantly after landfall.For example, over 150 years, northeastern parts ofthe peninsula experienced a minimum of one F3 hur-ricane, six F2 hurricanes, and thirty F1 hurricanes,while southwestern parts experienced no F2 or F3damage and fewer than five F1 storms. Though a sig-nificant disturbance across much of the Yucatan, hur-ricanes may have shorter-lived and less severe eco-logical impacts than fire or human land use. Howeverthe interaction of these factors (e.g., fires followinghurricanes) may be significant and deserves furtherstudy.

Forest Response to an Experimental Hurricane

The hurricane experiment simulates the forestimpact of an intense storm like the 1938 Hurricane.In October 1990 canopy trees were pulled over usinga winch, based on data from 1938, resulting in directand indirect damage to nearly 70 percent of thestand. Vegetation response has been studied over tenyears and this year all new trees greater than 5 cmdiameter were measured by Audrey Barker Plotkin.Most saplings are advance regeneration that waspresent in the understory of the stand before themanipulation. Regeneration is dominated by blackbirch, followed by red maple and yellow birch. Red

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Smoothed regional gradients in reconstructed hurricanedamage across the Yucatan Peninsula, Mexico, showingaverage return intervals at different damage levels(1851–2000). Damage ranges from F0 (loss of leaves andsmall branches) to F3 (most trees blown down).

Summer student Kristin Wilson and AudreyBarker Plotkin.

oak was dominant in the pre-manipulation forest, butonly one red oak has reached the 5-cm size class since1990. This compositional shift is dampened some-what by survival of some of the original canopy,which currently comprise more than one-third oftrees ≥ 5 cm diameter. These survivors include pros-trate stems, bent and leaning trees that are rebuildingtheir crowns, and vigorous undamaged trees. Thelargest trees are undamaged red oaks (46 stems/ha),which dominate the area’s basal area. Smaller redmaples have persisted as well. The current foreststand has a multi-layered structure, with densesaplings forming a lower stratum below a middlelayer of recovering trees, and scattered tall emergentstems from the original stand.

The manipulation exerted a large effect on theunderstory as increased light reached the forest floorfor several years, uprooting of trees resulted in patch-es of disturbed soil, and pits and mounds covered 8percent of the area. Although overall floristic changewas slight, new species were especially common inplots containing a pit or mound. Two exotic andpotentially invasive species (Morrow’s honeysuckleand Oriental bittersweet) have been found in onepart of the experiment; this area will be monitoredclosely but they are not expected to increase due tothe dense forest conditions. Raspberry speciesappeared in the experimental site, probably fromburied seed but are now declining. Hay-scented fern(Dennstaedtia punctilobula) and starflower (Trientalis bore-alis) increased markedly after the manipulation, butare now declining, indicating that understory condi-tions may be returning to pre-disturbance conditions.However, two abundant shrub species, shadbush(Amelanchier spp.) and hazel (Corylus cornuta) continueto increase. The landscape context of the site (sur-rounded by intact forest) and rapid recovery of theforest canopy via advance regeneration and sprout-ing contribute to the observed stability.

Thirty-two Years of Change in a New England Forest

Permanent plots allow the long-term study ofchanges in forest composition and structure; howev-er, few datasets extend beyond thirty years. In 1969,soil scientist Walter Lyford established a a seven acrepermanent grid on the Prospect Hill tract in mixedhardwood forest that originated after an old-fieldwhite pine stand was cut in about 1890. The site waschosen as typical of many upland forests and Lyfordmapped the locations and sizes of all trees, downed

wood, tip-up mounds, human artifacts (e.g.,stonewalls, prior experiments), and soils. With resur-veys on approximately ten-year intervals the areaprovides an opportunity to track forest dynamics ona fine spatial scale, and to relate vegetation to envi-ronmental variation and past disturbance. AudreyBarker Plotkin is sampling the area to complete athirty-two year spatial database of change.

Mice and Songbirds: Vertical Distributions andPredator-Prey Interactions in Oak Forests

For the past four summers, Cathy Langtimm andBecky Field have studied songbirds and white-footedmice (Peromyscus leucopus) in forests on Prospect Hilland in Tom Swamp addressing the possibility thatmice are important predators on the eggs of open-nesting songbirds. White-footed mice are foundthroughout New England forests with both positiveand negative effects on the mixed conifer-deciduousecosystem. They may help to control insect infesta-tions by eating many insects including the pupae ofgypsy moths. They also eat seeds, which can eitherassist with regeneration from buried seed caches orreduce regeneration of plant species. Mice are alsopart of the cycle of transmitting Lyme disease; infect-ed mice can pass on the bacteria to deer ticks.

A little studied aspect of these mice is their climb-ing behavior. During this study, small mammals werelive-trapped on the ground, in shrubs, and in thecrowns of canopy oak trees. All years have shownmarked differences in population densities andweather conditions. Mice were captured in canopyoak trees, although at low frequencies. Climbingactivity in the understory increased from summerinto fall as the number of insects, fruit, and seeds onplants increased.

Nest predation is a major mortality factor inopen-nesting songbirds. Yet, most studies on nest suc-cess have looked at predation in ground or shrubnests. Important data are missing for nest predationand types of predation above shrub height. We havecompared relative nest predation rates in ground,shrub, lower canopy, and higher canopy levels, usingartificial nests baited with plasticine eggs set out inMay, June, and July. In the summer of 2000, thiswork continued with the help of Scott Demers, asummer student supported by Harvard Forest andthe Department of Natural Resource Conservation,University of Massachusetts. Preliminary analysisshowed that annual predation rates appeared to fluc-

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tuate with changes in rainfall during the summers.“Bird predation,” defined by complete removal ofboth eggs, appeared to be higher in the upper strataand in May trials. “Small mammal predation” wasidentified by chewed plasticine eggs, many of whichhad tooth marks from white-footed mice. This typeof predation tended to be higher at lower heightsalthough still evident in the higher strata. Thesetrends suggest that there are differences in predationat different heights and that small mammals arecapable of climbing into the canopy and preyingupon songbird nests.

Acorns are important for winter survival of miceand estimates of acorn abundance at both trappingsites continued in a collaborative study with JohnO’Keefe. Estimates for fall 1998 and 1999 successfully predicted mouse population numbers the fol-lowing years.

Botanical Studies

Barry Tomlinson continued work on Calamus, the rattan cane of commerce, to show how the unusualvascular anatomy originates in the developing shoot.These results clearly show that vascular bundles areinitiated long before the leaves they supply. It wasestablished that the unusual gymnosperm Gnetum hasreaction fibers (gelatinous fibers) like those in woodyangiosperms. However, the fibers are extra-xylaryand play an important role in establishing and main-taining the distinctive architecture of this plant.

Graduate Student Research

Rebecca Anderson completed fieldwork for her MFSstudy of the role of climate and succession on thedevelopment and lateral expansion of forested peat-lands in New England. Work on three peatlandsinvolved detailed mapping of the basin morphome-try, characterization of the underlying sediments, andradiocarbon dating of major changes in develop-ment. The sites have similar histories, evolving froman open lake to sedge and shrub wetlands and thento swamp forest that spread from the original basinonto the surrounding upland. While sediment sam-ples are being dated, Rebecca is developing a modelfor forest peatland development and is beginning tointerpret the influence of climate on this history.

Jesse Bellemare continued his thesis researchinvestigating the impacts of past land-use on richmesic forests (RMF) in western Massachusetts. RMFare uncommon communities dominated by sugarmaple and white ash, with a lush herb layer includingspring ephemerals and unusual fern species. Thecommunity has high conservation value due to thediversity of uncommon species. The potential sensi-tivity of woodland herbs to past human disturbance,particularly nineteenth century forest clearance foragriculture, has led researchers to speculate that thecomposition of RMF may be strongly linked to pastland use. However, the absence of detailed historicalrecords has precluded an accurate assessment ofthese effects in much of the eastern United States.Jesse is using the 1830 land use, together with twen-tieth century data, to reconstruct forest cover over thepast two centuries. Vegetation sampling on RMFsites with differing histories is documenting variedspecies responses to past land use. These responsesappear to result from species-specific life history

14

P. Barry Tomlinson, E. C. Jeffrey Professor of Biology, Emeritus.

traits, including characteristics such as seed production rate and dispersal mode. Certain ant-dispersed and gravity-dispersed species are stronglyassociated with continually forested sites (primaryforests), whereas bird-dispersed and wind-dispersedspecies tend to be present in these as well as in sitesthat were historically cleared for agriculture (second-ary forests).

A striking illustration of this pattern can be seenin the contrasting distributions of three-leaved tooth-wort (Dentaria maxima), a clonal species which pro-duces little or no seed, and baneberry (Actaea rubraand A. pachypoda), which produces abundant bird-dis-persed fruits. Three-leaved toothwort is common inprimary RMF, but rarely occurs in secondary stands,likely due to seed limitation. In contrast, baneberrycan successfully disperse its berries by birds andmammals into secondary forests. One striking excep-tion to this trend is provided by bloodroot,(Sanguinaria canadensis), an ant-dispersed species typi-cal of RMF, which is more abundant in secondaryforest than in primary. This unusual distribution pat-tern is apparently due to bloodroot’s ability to persistand thrive in the high-light environment ofhedgerows, from which it has successfully recolo-nized secondary stands as they develop. This studystresses the varied responses of species to past distur-bance, underscores the legacies of land use, and high-lights the importance of considering history whenevaluating the factors influencing plant communitycomposition and species distributions.

Rob Eberhardt completed his MFS thesis on theinfluences of land-use history, fire, and environmen-tal conditions on patterns of forest vegetation onCape Cod National Seashore and submitted a manu-script for publication. He is continuing to work onanalyses for the entire Cape with Glenn Motzkin andDavid Foster.

Matt Kizlinski completed his field and laboratorywork for his MFS thesis examining vegetation andecosystem responses following hemlock logging. Inresponse to the expanding infestation of hemlockwoolly adelgid (HWA), extensive hemlock forests arebeing logged, often before infestation. Since hemlockhas not been a valuable timber species and has beenlargely ignored by silvicultural research, the long-term fate of these logged sites is unknown. To betterunderstand recovery from logging, ten sites rangingfrom one to thirteen years of post-logging successionwere chosen for intensive vegetation and ecosystemstudy. Vegetation quickly establishes in the high-lightenvironment created by logging. Unlike the slower,less uniform vegetation response accompanyingHWA infestation, nearly the entire harvested arearesponds with vegetation regrowth. Species not nor-mally found in healthy hemlock forests become com-mon, such as thickets of blackberry and raspberry.Within a few years, black birch seedlings dominateand grow to form a low dense canopy, shading outthe early successional species.

Ecosystem changes following hemlock loggingare equally dramatic. The thick soil layer of hemlock

15

Typical rich mesic forest plant species showing differering distribution patterns in response to past land use. Three-leavedtoothwort (Dentaria maxima) (a) is a vegetatively-reproducing species that rarely produces seed; it occurs commonly inprimary forest, but is absent from secondary stands. White baneberry (Actaea pachypoda) (b) and red baneberry (A. rubra)produce seeds abundantly in the form of bird-dispersed berries; they occur commonly in both primary and secondarystands. Bloodroot (Sanguinaria canadensis) (c) produces seeds that are ant-dispersed, suggesting that the species could expe-rience difficulty recolonizing secondary forest. However, bloodroot has persisted and thrived throughout the agriculturallandscape in hedgerows (d), from which it has successfully recolonized many secondary stands.

needles is gradually replaced by a thin layer of decid-uous leaves. Despite this large loss in total forest floormass, old cuts maintain total nitrogen mineralizationrates that are similar to intact hemlock stands.However, a much larger proportion of the nitrogen isin the form of nitrate, which is uncommon in intactecosystems in the northeast. Higher nitrification ratescan also be seen in younger cuts; due to a lack ofplant uptake, nitrate could possibly be leaching out ofthe system. This project links previous and ongoingresearch at the Harvard Forest. Studies of the directimpact of HWA on ecosystem properties, combinedwith HWA’s indirect effect of increased hemlock log-ging, highlight the widespread and severe impact thisinsect’s introduction is having on hemlock forests.

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. Eachsite addresses questions of a long-term nature; collec-tively the sites undertake comparative studies acrossecosystems. Representatives from each site and NSFmeet twice annually to coordinate network-wideactivities and to collaborate. The central theme of theHarvard Forest LTER is interpretation of the struc-ture, composition, and function of forest ecosystemsin terms of their history of natural and human dis-turbance and environmental change. This research isbeing addressed at the stand, landscape, sub-region(e.g., central Massachusetts), and regional (NewEngland) 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), Earth and Planetary Sciences (S.Wofsy, B. Munger), and Harvard Forest (D. Foster,D. Kittredge, G. Motzkin, D. Orwig) at HarvardUniversity as well as the Ecosystems Center-MBL,Woods Hole (J. Melillo, K. Nadelhoffer, P. Steudler),the Complex Systems Research Center at theUniversity of New Hampshire (J. Aber) and theUniversity of Massachusetts (M. Mulholland).Emery Boose is the LTER Data Manager with assis-tance from Julie Pallant. The research is organized tomaximize the interactions among scientists from dif-ferent disciplines. Four major scientific approachesinclude: (1) retrospective studies of historical changesin the environment and ecosystems; (2) long-term

measurements of forest structure and function; (3)experimental manipulations; and (4) synthesis andmodeling. The LTER science group meets approxi-mately monthly. The Harvard Forest EcologySymposium is held to present current research withabstracts published annually. The program for the2001 symposium 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.

Nitrogen Saturation in Temperate Forest Ecosystems

Human activity has augmented the cycling of nitro-gen through the biosphere more than any other

16

Heidi Lux on the soil warming plots.

element. As the pivotal element in amino acids, pro-teins, nucleic acid, chlorophyll and other biomole-cules, nitrogen has profound effects on ecosystemfunction. As a component of nitric acid, nitrogen is akey element in the acidification of soils and streamsdue to acid deposition.

In 1989, the LTER group from the University ofNew Hampshire, led by John Aber, published a seriesof hypotheses concerning the expected responses ofN-limited forest ecosystems to chronically elevated Ndeposition. Central to these was the expectation ofhighly non-linear or threshold responses in the induc-tion of nitrification and nitrate leaching. They alsopredicted that excess N deposition would eventuallyreduce tree growth and lead to forest decline.However, there was no real expectation that thiswould happen in a short period of time.

In 1998 John’s group revisited these initialhypotheses. Two of the most important ones havebeen supported by the findings. It is indeed possibleto induce nitrification and nitrate leaching withchronic N additions, although this took much longerin the very N-poor hardwood stand than we expect-ed. Mechanisms by which large amounts of added Nare retained in these systems are now a key area ofstudy. In addition, and rather surprisingly, the appli-cation of nitrogen did induce significant reductions ingrowth rates in the pine stand. By the year 2001, thepine stand receiving the highest level of N depositionwas effectively dead. To date there has not been any

growth decline in deciduous, broad-leaved stands.This result may assist us in interpreting historicalepisodes of forest decline and anticipating futuretrends in forest health in areas of high N deposition.

Carbon Sequestration by Temperate Forests

Northern mid-latitude forests appear to be importantsinks for as much as 25 percent of the CO

2added to

the atmosphere by fossil fuel burning since 1980. Thenet sink varies inter-annually by a factor of two ormore, correlating with global-scale climate variations,and possibly accelerating in the 1990s. When theHarvard Forest LTER was initiated in 1989, thegroup from Harvard’s Department of Earth andPlanetary Sciences headed by Steve Wofsy set out todetermine whether typical New England forests werea net sink for CO

2, how large the sink was, and what

processes controlled it. Various factors have beensuggested: previous land use, fire and forest manage-ment are factors that are subject to deliberate modifi-cation; longer growing seasons due to climate warm-ing, fertilization by elevated CO

2and N deposition

are environmental influences not subject to directmanipulation.

Rates for Net Ecosystem Exchange (NEE) ofCO

2at Harvard Forest have been measured by

Steve’s research group each hour since 1990, usingthe eddy covariance technique, along with extensivebiometric measurements of species-specific changes

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Long-term results from the nitrogen saturation experiment document the impact of two different doses of nitrogen(low, high) on pine and hardwood stands. The pine stands exhibited the greatest response in terms of nitrate leaching from the soils (left) and reduction in wood net primary production (right).

in the C stored in live and dead wood. These long-term observations allow assessment of the accuracyof the C budget derived from hourly flux data, andquantitative analysis of the observed response of theecosystem to environmental variation for time scalesfrom instantaneous (hourly) to decadal.

The long-term record of NEE shows average netuptake of 2.0±0.4 Mg C ha-1yr-1, with inter-annualvariations exceeding 50 percent Cumulative uptakecompared well with biometric data that indicatedstorage of 1.6±0.4 Mg C ha-1 yr-1 over eight years, 60percent in live biomass and the balance in coarsewoody debris and soils. Very different processes con-trol carbon uptake on long and short time scales.Ecosystem responses to weather and climate (e.g.,variations in growing season length or cloudiness)regulated seasonal and annual fluctuations. Legaciesof prior disturbance and land use, especially standage and composition, dominated longer time scales.Thus short-term variations of NEE at HarvardForest reflect prompt forest response to environmen-tal influences, while inter-annual variations reflect

interactions of climate variations with ecosystemproperties on annual time scales, affecting tree mor-tality, respiration rates, length of the growing season,and available light. Since seasonal and annual climat-ic anomalies are often coherent over large spatialscales, the processes described here are significant inmediating inter-annual variations of the rate ofincrease of atmospheric CO

2. The long-term carbon

budget at Harvard Forest reflects the slowest-chang-ing ecosystem properties: stand age and composition,soil fertility, and coarse woody debris. The enormousareas occupied by mid-successional forests (30–100years old) in the U.S. have been cited as the majorfactor in present terrestrial uptake of carbon, forwhich these studies provide strong quantitative sup-port and mechanistic understanding.

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(Left): Cumulative net carbon exchange at Harvard Forest, the sum of hourly data for nine years; (right): Comparisonof annual sums of net carbon exchange with ecological data showing long-term changes in carbon pools. These dataestablish the accuracy of long-term eddy flux measurements using conventional biometry, provided that averaging timesare sufficient (~five to eight years).

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Harvard Forest Ecology Symposium 2001 — Abstracts and Presentations

Anderson, R. Development and lateral expansion of peatlands in central New England.Barford, C. et al. Eddy covariance and biometric measurements of CO

2exchange at the Harvard Forest.

Barker Plotkin, A. and D. Foster. Long-term forest monitoring at Harvard Forest.Barker Plotkin, A. and D. Foster. Understory dynamics in the experimental hurricane.Barker Plotkin, A. , S. Martell, and D. Foster. Forest development after an experimental hurricane.Barnes, D. and S. Wofsy. Evaluation of greenhouse and ozone-depleting gases in rural New England.Bauer, G. et al. Is increased nitrogen availability predictive for long-term forest carbon sequestration?Bellemare, J., G. Motzkin, and D. Foster. History and species composition of rich mesic forests.Boose, E. The new Harvard Forest meteorological station.Borken, W., E. Davidson, K. Savage, and K. Angeloni. Response of CO

2release to dry/wet cycles.

Burk, J., G. Motzkin and D. Foster. The historical landscape of southern New England in early records.Cobb, R., D. Orwig, and S. Currie. Effects of hemlock woolly adelgid on foliar decomposition. Colman, B., K. Nadelhoffer, and W. Currie. Decadal scale recovery of 15n tracers in bolewood.Currie, W., K. Nadelhoffer, and B. Colman. Woody detritus, land use and long-term C and N interactions.Dail, D., E. Davidson, J. Chorover, L. Dria, and P. Hatcher. Rapid abiotic transformation of nitrate in soils.Davidson, E., K. Savage, and S. Trumbore. Interannual variation of CO

2production in the O horizon.

Dria, K. et al. Changes observed by NMR and pyrolysis gc/ms of Harvard Forest soils.Eberhardt, R. et al. Land-use legacies in the vegetation of Cape Cod national seashore.Fitzjarrald, D. and K. Moore et al. Forest-atmosphere exchange processes: report on activities 2000-2001.Francis, D. et al. Climate change in New England inferred from Chironomid remains in lake sediments. Fuller, J. et al. Recent climate reconstruction in New England forests.Hadley, J. and J. Schedlbauer. Ecosystem carbon exchange in an old-growth hemlock forest.Horii, C., J. Munger, S. Wofsy, and M. Zahniser. Atmospheric reactive N partitioning and fluxes.Kaufman, S. and F. Bazzaz. Effects of n deposition on Alliaria petiolata and indigenous species.Kittredge, D. and A. Finley. North Quabbin region’s chapter 61 spatial database.Kittredge, D., A. Finley, and D. Foster. Pattern of timber harvest in a complex forest landscape.Kizlinski, M. et al. Ecosystem and vegetation response to hemlock logging.Köster, D. and R. Pienitz. Paleoecological and ecological studies on diatoms in four New England lakes. Lewis, K. and F. Bazzaz. Responses of invasive species to release from herbivory.Lux, H., J. Melillo, T. Ahrens, P. Steudler, and F. Bowles. Warming, C storage, and feedbacks to vegetation.Magill, A. and J. Aber. Chronic nitrogen additions to two forest stands.Micks, P. and L. Nadelhoffer. Forest soil respiration and organic matter dynamics.Min, Q. and K. Moore. Regional climatology of cloud and aerosol for forest-atmosphere exchange. Moore, K., D. Fitzjarrald, and Q. Min. Modeling the effect of diffuse light on canopy CO

2 uptake.

Munger, J., J. Field, J. Budney, and S. Wofsy. Peroxyacetylnitrate concentrations at Harvard Forest. Muth, C. and F. Bazzaz. Tree seedling canopy responses to conflicting photosensory cues.Muth, C., K. Capecelatro, and F. Bazzaz. Influence of branch processes on tree canopy displacement.O’Keefe, J. Regeneration following clearcutting of red pine overstory — year 11.O’Keefe, J. and S. Johnson. Woody species phenology, Prospect Hill tract, Harvard Forest — 2000.Orwig, D., D. Foster, and D. Mausel. Patterns of hemlock decline due to the hemlock woolly adelgid. Orwig, D., G. Motzkin and D. Foster. Vegetation and history of ridgetop pitch pine and red pine.Orwig, D., R. Cobb, M. Kizlinski, and D. Foster. Ecosystem analyses of hemlock woolly adelgid. Parshall, T. et al. Landscape-scale history of vegetation and fire on Cape Cod, Massachusetts.Parshall, T. Variability in the long-term occurrence of fire in New England.Syfert, M., G. Motzkin, and B. Hall. Massachusetts in 1830: woodlands and cultural events.Verheyen, L. and M. Hermy. A comparison of European and North American ancient forest plant species.

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, K. Moore), Woods Hole ResearchCenter (E. Davidson), University of California (S.Trumbore), U.S. Geological Survey (E. Sundquist)and Harvard Forest (J. Hadley).

BULLARD FELLOWS

Kathleen Donohue (University of Kentucky) con-ducted research on the population biology and evolu-tionary ecology of seed dispersal and germination.She established a large field experiment in Kentuckyand New England to compare the adaptive value andgenetic basis of germination behavior between thesesites. She also initiated studies of the quantitativegenetic basis of seed dispersal and its plasticity.Kathleen presented seminars at Cornell Universityand Utah State University and developed manu-scripts concerning the influence of dispersal on multi-level natural selection, the influence of germination onlife-history expression and evolution, the physiologi-cal basis of germination responses to seasonal cues,and natural selection in heterogeneous environments.

Mary Ann Fajvan (West Virginia University)analyzed the ten years of regeneration data from theTom Swamp Hurricane study. She developed modelsto describe the height growth response of key treespecies and is using spatial statistics to describe regeneration establishment patterns. With AudreyBarker Plotkin, she is preparing two manuscriptsfrom the analyses. She also was an invited speaker at the Pennsylvania State University Forestry IssuesConference on sustainable forest management practices.

Manuel Lerdau (State University of New York,Stony Brook) began two projects in the Departmentsof Earth and Planetary Sciences and Organismic andEvolutionary Biology: (1) development of a coupledtunable diode laser absorption spectrometer with acontrolled environment leaf chamber and infraredgas analyzers for the measurement of the physiologi-cal regulation of nitrogen dioxide exchange betweenplants and the atmosphere; and (2) examination ofthe role of phylogeny and phenology in regulatingleaf chemistry, decomposition, and nutrient release inthe tropical deciduous forest of Costa Rica.

Lucinda McWeeney (Yale University) completedfour manuscripts on archaeology and environmentalreconstruction in New England and processed north-ern New Hampshire sediment cores for macrofossilsand pollen and archaeological sediments for charcoaland phytoliths. Lucinda participated in a regionalpaleoenvironment workshop at the PeabodyMuseum, Andover and presented seminars at theSociety for American Archaeology Annual Meetingin New Orleans. She traveled to South Carolina andVirginia to gather sediment samples for future paleo-ecology projects relating to pre-Clovis habitation sitesin the southeast.

Fraser Mitchell (Trinity College, Dublin) investi-gated long-term stand dynamics in relation to distur-bance regimes in temperate forests in New England.He investigated and attempted to quantify the impactof human versus natural disturbance to the forest atthe Pisgah old-growth stand in southern NewHampshire, a site at higher altitude than theMassachusetts sites, but more significantly, sufferingminimal impact from European settlement. Results ofpollen analysis from two sites at Pisgah have revealedthat forest composition today is remarkably similar tothat of the pre-European settlement (and pre-immi-gration of chestnut) forest. The most striking differ-ence between Pisgah and the Massachusetts sites isthat beech has remained as an important componentat Pisgah while it has declined following Europeansettlement in Massachusetts. More intensive dataanalysis is planned to quantify this difference. Theseresults raise questions about the tolerance of beech todisturbance and in particular its ability to recolonize.This issue is particularly relevant to the potentialimpact of fungal pathogens on the long-term survivalof beech in New England. Fraser completed pollenanalysis on samples from a site in Scotland. He inter-acted with staff members in the area of charcoalanalysis. This is a area of paleoecology that has been

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under-investigated in Ireland but certainly worthy ofattention in the future.

Karn Deo Singh (Harvard University) preparedguidelines for assessment of forest biological diversi-ty and phase one of a research proposal integratingbiological diversity conservation in forest manage-ment planning in Malaysia, which is currently beingimplemented by specialists from the Forest ResearchInstitute of Malaysia and Harvard University.

Fred Swanson (U.S. Forest Service, PacificNorthwest Research Station) worked on the AndrewsForest LTER synthesis volume and manuscripts con-cerning ecosystem response to the 1980 eruptions ofMount St. Helens, road ecology, and the dynamics ofwood in rivers. Collaboration with David Foster on apaper concerning LTER studies of land-use legaciesdrew significantly on their perspectives from cultural,managed, and wild landscapes of New England andthe Pacific Northwest. He gave seminars and classlectures at Harvard, University of Massachusetts,University of Vermont, Cornell, and the Institute ofEcosystem Studies.

Bullard Fellows for 2001–2002 include AlisonBerry (University of California, Davis), ElizabethColburn (Massachusetts Audubon Society), AaronEllison (Mt. Holyoke College), Colin Orians (Tufts

University), Akile Pite (Scientific Commission inForestry, Albania), Kimberly Smith (University ofArkansas) and Navjot Sodhi (National University ofSingapore).

EDUCATIONAL ACTIVITIES

Barry Tomlinson taught Harvard courses, Biosciences51, “Integrative Biology of Organisms” in the fallsemester and Biosciences 24, “Biology of Plants” dur-ing the spring semester. He also conducted a work-shop in “Tropical Plant Biology” at the FairchildTropical Garden, Miami, Florida, during summer2000. David Foster taught Biology 160, “ForestEcology” and Biology 199r, “Forest Practice andResearch.” Barry, David, John O’Keefe and other staffmembers led the Harvard Forest seminar of elevenstudents in the spring. David served on thesis com-mittees of three undergraduate students at Harvardand graduate students at the University of Massa-chusetts, University of Maine, and Laval University.Glenn Motzkin, Dave Orwig, and David supervisedthe four students in the Masters of Forest Science pro-gram. Dave Kittredge and David Foster worked close-ly with former Harvard undergraduate Mary Berlikto revise her thesis into a Harvard Forest Paper.

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David Kittredgeand Andy Finley.

Summer Research Program

The Harvard Forest Summer Student Research pro-gram, coordinated by Edythe Ellin and assisted bySarah Laubscher, attracted a diverse group of stu-dents to receive training in scientific investigations,and experience in long-term ecological research.Students work closely with researchers and manyconduct their own independent studies. The programincludes weekly seminars from 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 theAnnual Summer Student Research Symposium.

Summer Students 2001

Christian Arabia Allegheny CollegeTricia Burgoyne University of WisconsinPeter Cowan Kalamazoo CollegeBridgid Curry University of Notre DameCaitlin Dwyer-Huppert Clark UniversitySusan Italiano Gettysburg CollegeLynda Joudrey University of New

HampshireSpencer Meyer Dartmouth CollegeRosa Navarro Stanford UniversityJulia Silvis Harvard UniversityKathleen Theoharides Dartmouth CollegeMorgan Tingley Harvard UniversityLinda Jane Wan University of

PennsylvaniaKristin Wilson Middlebury CollegeTruus Thomas Benedict College

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Front (left to right): Sarah Laubscher (Proctor), Spencer Meyer, Rosa Navarro. Middle (left to right): SusanItaliano, Truus Thomas, Linda Jane Wan, Bridgid Curry. Back (left to right): Christian Arabia, CaitlinDwyer-Huppert, Kathleen Theoharides, Kristin Wilson, Julia Silvis. (Missing from photo: Tricia Burgoyne,Peter Cowan, Lynda Joudrey, and Morgan Tingley.)

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 ecology, conserva-tion, and management. The Museum also provides aunique setting for conferences and workshops spon-sored by the Forest and outside organizations. Dr.John O’Keefe has primary responsibility for thedevelopment of activities and coordination of the useof the Museum.

The Museum enjoyed another very successfulweekend schedule last year, welcoming over 1,100visitors on Saturday and Sunday afternoons, Maythrough October. Mary Ann Walker again receivedspecial thanks for her continuing, enthusiastic workas volunteer coordinator. Bill and Marianna Berry,Hector Cameron, Walt Davidson and MarthaSiccardi received special recognition for being themost active volunteers during the season. On a moresolemn note, this past spring the group was deeplysaddened by the death of Helen Gronich. who servedas volunteer coordinator for our first six years.

In late April Teresa Jones assumed the part-timeposition as Assistant Schoolyard LTER coordinator,responsible for coordinating Harvard Forest’s collab-oration in ecological studies with schools in theregion. Teresa is updating the interpretation on ournature trails and will develop new a interpretation forthe boardwalk through Black Gum Swamp.

During the year the Museum provided programsfor twenty-four elementary and secondary schools,forty-three college and university classes, and twenty-three community and professional groups. InSeptember the Forest hosted a group of ChineseLTER scientists touring selected North AmericanLTER sites to learn about our research programs. InNovember more than forty silviculture professors vis-ited the Forest as part of a field trip in conjunctionwith the National Society of American Foresters con-vention in Washington, D.C., and in June the Forestand Museum hosted a field trip in conjunction withthe American Geophysical Union meeting in Boston.

Meetings, Seminars, Conferences

The Twelfth Annual Harvard Forest Long TermEcological Research Symposium was held in theMuseum on April 23 followed by the NationalInstitute for Global Environmental Change meetingon April 24. Other meetings at Harvard Forest

included meetings of the North Quabbin RegionalLandscape Partnership, Massachusetts WatershedCoalition, Massachusetts Extension Service CovertsProject, Massachusetts Department of Environ-mental Management Logging Safety Workshop,Massachusetts Land Trust Coalition, MassachusettsAssociation of Professional Foresters, MassachusettsExecutive Office of Environmental Affairs, MillersRiver Watershed Council, Natural Resources Con-servation Service, New England Chapter of the Wild-life Society, Northeastern Loggers Association,Predator Conservation Alliance, Project LearningTree, Vegetation Control Service, Athol Bird andNature Club, and the Nature Conservancy. TheForest hosted faculty retreats for the University ofMassachusetts Department of Natural ResourceConservation and Harvard’s Department of Organ-ismic and Evolutionary Biology.

Speakers in the Harvard Forest Seminar series:

Carl Beierkuhnlein University of Rostock, Germany

Dick Boisvert Deputy State Archaeologist for New Hampshire

Julie Brigham-Grette University of Massachusetts Kim Brosofske University of Rhode IslandDena Dincauze University of MassachusettsKathleen Donohue University of KentuckyMary Ann Fajvan West Virginia UniversityAdrien Finzi Boston UniversityRichard Forman Harvard University Megan Griffiths Tufts UniversityOlivier Honnay Catholic University, BelgiumMalcolm Hughes University of ArizonaBrian Jones Mashantucket Pequot

MuseumManuel Lerdau State Univ. of New York Glenn Matlack University of S. MississippiLucinda McWeeney Yale University Wyatt Oswald University of WashingtonStéphanie Pellerin Université LavalNathan Phillips Boston UniversityMichael Reed Tufts UniversityBryan Schuman Brown UniversityK. D. Singh Harvard UniversityPeter Siver Connecticut CollegeFred Swanson H. J. Andrews Experimental

Forest Steven Trombulak Middlebury CollegeKris Verheyen Catholic Univ., Belgium

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FOREST MANAGEMENT AND MAINTENANCE

After more than twenty years of service to HarvardForest, Charles “Pete” Spooner retired from thewoods crew in January. The contributions Pete hasmade to the Forest are manifold and can be seen bestin the local landscapes he has helped to shape andchange. Pete has been a mainstay here, always thereto lend a hand or supply advice. Many technicalproblems in research and equipment have beensolved with Pete’s expertise and his knowledge of thelocal countryside. His welding and steel fabricationskills are evident throughout the physical plant. Weare fortunate to be able to rely on Pete for part-timeassistance in the future.

This past spring we completed the final land-scaping details on the courtyard connecting ShalerHall, Torrey Lab, and the Archives. These improve-ments have received many positive comments andinspired us to continue to the front of Shaler Hallwhere we are widening the parking lot and addingnew lighting, walkways, paving, and granite edging.This effort is part of a three-phase project to improveaccess and parking for growing numbers of visitorsand staff.

Other efforts include the clearing and fencing forpasture of a new field on the west slope below Fisher

House, the installation of our new weather station,permanent 110 volt AC power to the new canopytower in the hemlock stand, as well as continued thin-ning and improvement cutting in the softwood plan-tation complex at the western edge of Harvard Pond.

LONG TERM DATA MANAGEMENTHarvard Forest Archives

The Archives, which represents the map, document,photographic, correspondence, and research samplecenter of our data management system, serves as adaily resource for Harvard Forest researchers andvisitors from many colleges and historical, ecological,and conservation organizations. As part of the effortto develop a management plan for the HarvardForest properties, historical deed, map, and corre-spondence files were researched for land-use provi-sions and historical information. A summary of allproperty transactions from 1907 to the present wasexpanded and a simplified property and deeds filewas created, combining documents from manysources. Several hundred building and highway planswere also added to the map files. Other additions tothe map collection included comprehensive coverageof the nineteenth century Coast and Geodetic

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Charles “Pete” Spooner and David Foster discussing the renovation of the front of Shaler Hall.

surveys for the coastal region from New York toCape Cod, and twentieth century U.S.G.S. topo-graphic quadrangles. In conjunction with theMassachusetts 1830s mapping project, the entireseries was acquired from the state archives, givingHarvard Forest a complete set of this unique andvaluable resource. To create a central archive data-base, files documenting maps, samples, aerial photos,research files, and publications were inventoried,updated, and converted to the ProCite bibliographicformat.

Historical and current slides and photographswere used to create a detailed photo gallery on theHarvard Forest Web site. Lantern slides were sortedand cataloged, with several high-quality prints pro-duced from the glass originals. A large series of pho-tographs of Cape Cod and the Islands was added tothe aerial photo collection and many historical pho-tos were duplicated or scanned for use in projects.Records for the Sanderson family, which farmed por-tions of Prospect Hill in the eighteenth and nine-teenth centuries, were strengthened by acquisition ofthe family’s original account book from a localbookdealer, providing a primary account of agricul-tural activity on the tract. In addition, Kathleen

Hunter, a Sanderson descendant, made severalresearch visits and donated a copy of the familygenealogy records to the archives.

Approximately 70 boxes of new material wereadded to the sample archive, including soils from thecoastal, soil warming, and adelgid studies, and treecookies from the ancient Gribben forest in Michigan.

Computers

Major improvements to our computer facilities con-tinued this year with funding from NSF and internalsources. The network was extended to the Fisher,Raup, and Community Houses (via optical fiber) andto the EMS Tower (via DSL modems). FourWindows 2000 servers, a high-capacity tape drive,and a CD-Tower were purchased, as well as threenew staff computers. New graphics equipmentincluded a portable computer projector and laptopcomputer, digital camera, and a large-format opticalscanner. A second computer projector will be includ-ed in the remodeled Seminar Room. Several long-term bibliographic databases were consolidated andconverted to a common format (ProCite) forimproved access and maintenance.

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(Left): Emery Boose and the new meteorological station,(right): The old met station.

ACTIVITIES OF THE HARVARD FOREST STAFF

Rebecca Anderson presented a poster at the LTERAll Scientist’s Meeting (ASM) in Snowbird, Utah.She led field trips to local peatlands for classes fromthe Harvard School of Landscape Architecture,Harvard College, and Mount Holyoke College and atour of Harvard Forest’s Black Gum Swamp for stateBiodiversity Days. Audrey Barker Plotkin presenteda poster at the LTER ASM, attended the ForestStewards Guild Annual Meeting in Silver Bay, NewYork, and presented a seminar, “Forest Response toSimulated Hurricane Damage,” at the University ofMassachusetts, Boston.

John Burk participated in conservation meetingsand conferences, including the inaugural MountHolyoke “Summit on the Range” for the ConnecticutValley, and led a tour of the Quabbin Reservoir forREU students and staff. He served as Petershamcoordinator for the Massachusetts Biodiversity week-end.

Susan Clayden attended the Eighth InternationalPaleolimnology Symposium at Queen’s University,Kingston, Ontario.

Richard Cobb presented talks at the EcologicalSociety of America (ESA) meeting and at theUniversity of Maine Cooperative ExtensionWorkshop for practicing foresters.

David Foster presented lectures at the LandscapeLegacies of Human Land Use symposium at Duke, Ecotarium (Worcester Natural HistorySociety), and Sturbridge Village, based on LTERresearch, Thoreau’s Country, and New England ForestsThrough Time. He joined Eric Chivian and PaulEpstein from Harvard’s Program for Human Healthand the Global Environment at Aerlie House inVirginia for a two-day briefing of congressional aideson environmental issues, where he discussed issuesrelating to fire ecology and management. David rep-resented HF at LTER Coordinating Committeemeetings and was interviewed by Jamie Sayen forWild Earth and Northern Forest Forum. David con-tinued in his capacity as editor for Ecosystems,Progress in Physical Geography, and NortheasternNaturalist, board member for the ConservationResearch Foundation, advisor to the HighsteadArboretum, and became a trustee of the Trustees ofReservations. At Harvard he serves as the Director ofGraduate Studies for the M.F.S. program, an execu-tive committee member for the NIGEC program,and served on a faculty search committee for OEB.

He travelled with a Harvard Museums of NaturalHistory (HMNH) group to Patagonia, and with theAlumni Association and HMNH to the GalapagosIslands.

Donna Francis presented a seminar at theGeological Society of America, Northeast Section, inBurlington, Vermont.

Julian Hadley presented talks at ESA and attend-ed the annual Ameriflux meeting in Atlanta where hepresented the first direct measurements of carbonexchange in an eastern hemlock forest.

Brian Hall continued as the weather stationobserver, adding to eighty-eight years of weatherrecords at Harvard Forest. This year we are collect-ing data at the old, manual site and at the new, auto-mated meteorological station concurrently for oneyear to allow evaluation of data compatibility.

Dave Kittredge served as invited keynote speak-er on forest policy at Penn State’s Forest Issues con-ference. The Ford Foundation awarded Dave aresearch fellowship to study forms of private ownercooperation in temperate forest systems. He will usethis opportunity to spend his upcoming sabbaticalyear at the Forest pursuing these ideas, and examplesfrom around the world.

Matt Kizlinski presented his research at a semi-nar at the Arnold Arboretum.

Glenn Motzkin presented a poster on coastalresearch at ESA and attended the Long Island PineBarrens Research Forum at Brookhaven NationalLaboratory. He led a field trip through theConnecticut Valley for David Foster’s Forest Ecologyclass and a trip to Montague Plain for the ConwaySchool of Landscape Design. Glenn served on thegraduate committees of Rebecca Anderson, JesseBellemare, and Rob Eberhardt at Harvard Forest andSally Shaw from the University of Massachusetts.Glenn was featured in an interview on forestresilience and recovery in the Woodland Steward. Hecontinued to serve as an Ecology Advisor for TheTrustees of Reservations and was appointed as anAssociate Member of the Massachusetts NaturalHeritage and Endangered Species Program AdvisoryCommittee.

Sylvia Barry Musielewicz received her Master’sdegree from the University of Minnesota and pre-sented a poster on her work in Malawi at a meetingof the Geological Society of America.

John O’Keefe gave talks on the history of north-eastern forests at the University of Massachusetts,Fruitlands Museums, Shelburne Farms in Vermont in

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conjunction with the National Park ServiceConservation Institute, SUNY, Albany, MountHolyoke College’s Sense of Place lectures, the NewEngland Museum Association meeting in Portland,Maine, and the New England Society of AmericanForesters’ Advances in Forest Biology Workshop. Herepresented Harvard Forest on the ENFOR DistanceLearning in Forestry planning group, cosponsored bythe Lincoln Institute of Land Policy and the KennedySchool of Government. John attended the LTERASM and ESA in Snowbird, Utah, presented a posteron the long-term impacts of land use history at the joint American Society for EnvironmentalHistory/Forest History Society meeting in Durham,N.C. and attended the NSF LTER CoordinatingCommittee meeting in Phoenix. He was a judge atthe Mahar High School Science Fair, and serves onthe boards of the Millers River Watershed Counciland Mount Grace Land Conservation Trust, wherehe is currently vice president, and on the executivecommittee of the North Quabbin RegionalLandscape Partnership. He also continues to serveon the Quabbin Science and Technical AdvisoryCommittee and Secretary Durand’s Advisory Groupon Environmental Education.

Dave Orwig attended the LTER ASM and presented a talk at ESA. He presented hemlock woolly adelgid (HWA) research findings at a publichearing organized by State Senator Brewer at MountWachusett Community College, at the NorthAmerican Forest Insect Work Conference in Ed-monton, Canada, and at the Massachusetts ForestryAssociation Annual meeting. Dave helped lead a fieldtour at Harvard Forest to the Visiting SilvicultureInstructors tour. In addition, he gave a seminar onold-growth forests and led a trip to Mount Wachusettfor the Massachusetts Audubon Society. He wasappointed Chairman of the Silvicultural Recom-mendations section of the U.S.D.A. HWA StrategicManagement Plan and serves on Matt Kizlinski’sM.F.S. committee.

Tim Parshall gave talks at the AmericanGeophysical Union and the Pine Barrens Symposiumin Brookhaven, N.Y. and presented research results at the ESA. He participated in a workshop for the Massachusetts Department of EnvironmentalProtection on sediment coring protocol for deter-mining mercury level in Massachusetts lakes.

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David Foster.

John O’Keefe.

John Aber University of New HampshireToby Ahrens Ecosystems Center — MBLMark Ashton Yale UniversityWilliam Bain Harvard UniversityPeter Bakwin Harvard UniversityCarol Barford University of WisconsinDiana Barnes Harvard UniversityFakhri Bazzaz Harvard UniversityDennis Balocchi NOAA/ARL/Oak Ridge, Tenn. K. Boering Harvard UniversityRichard Bowden Allegheny College Frank Bowles Ecosystems Center — MBLAlfram Bright Harvard UniversitySean Burrows University of WisconsinJeannine C-Bares Harvard UniversityMichael Canfield Harvard UniversityChaur-Fong Chen Oregon State UniversityAlan Coleman Harvard UniversityWilliam Currie Virginia Polytechnic Univ.David B. Dail University of GeorgiaEric Davidson Woods Hole Research CenterBruce Daube Harvard UniversityFrederik Debaeke Kuleuven, BelgiumMarty Downs Ecosystems Center — MBLTodd Drummey Ecosystems Center — MBLJim Ehleringer University of UtahBob Evans U.S.D.A. Forest ServiceRebecca Field University of MassachusettsDavid Fitzjarrald SUNY, AlbanySon-Maio Fan Harvard UniversityAndrew 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 ServiceDaniel Jacob Harvard UniversityChristine Jones Harvard UniversitySylvia Kaufman Harvard Unversity

Chris Kerfoot Ecosystems Center — MBLOtto Klemm Univ. of New HampshireJuri Knjazikhin Boston UniversityTakashi Kohyama Hokkaido UniversityChun-Ta Lai University of UtahCathy Langtimm U.S.G.S.; Holy Cross CollegeBarry Lefer Univ. of New HampshireHeidi Lux Ecosystems Center — MBLAlison Magill Univ. of New HampshireEls Malfait Kuleuven, BelgiumLynn Margulis University of MassachusettsMary Martin Univ. of New HampshireJerry Melillo Ecosystems Center — MBLJennie Moody University of VirginiaKathleen Moore SUNY, AlbanyJeff Morisette NASASarah Morisseau Ecosystems Center — MBLMitch Mulholland University of MassachusettsRanga Myneni Boston UniversityJ. William Munger Harvard UniversityChristine Muth Harvard UniversityKnute Nadelhoffer Ecosystems Center — MBLJeffrey Parker Smithsonian Environ. Res.Sophie Parker Ecosystems Center — MBL Ronald Prinn M.I.T.Elizabeth Pyle Harvard UniversityMichael Rogers GA Institute of TechnologyKathleen Savage Woods Hole Research CenterDavid Schaub University of ArizonaTim Sipe Franklin & Marshall CollegePaul Steudler Ecosystems Center — MBLBritt Stephens Harvard UniversityKristin 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, BelgiumLisa Windham Lehigh UniversityGreg Winston U.S. Geological Survey

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VISITING RESEARCH SCIENTISTS AT THE HARVARD FOREST 2000–2001

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

PUBLICATIONS

Aber, J. D. and R. Freuder. 2000. Sensitivity of a for-est production model to variation in solar radia-tion data sets for the Eastern U.S. Climate Research15: 33–43.

Barker, M. and D. Péréz-Salicrup. 2000. Com-parative water relations of mature mahogany(Swietenia macrophylla) trees with and withoutlianas in a subhumid, seasonally dry forest inBolivia. Tree Physiology 20: 1167–1174.

Barnes, D. H. 2000. Quantifying urban/industrialemissions of greenhouse and ozone-depletinggases based on atmospheric observations. Ph.D.Thesis, Harvard University.

Barry, S. L. 2001. Stratigraphic correlation andgeochronology of varved sediments from LakeMalawi, East Africa. Master’s Thesis, Universityof Minnesota.

Bazzaz, F. A. and S. Catovsky. 2001. Resource parti-tioning. Pp. 173–183 in S. Levin (Ed.), Encyc-lopedia of Biodiversity. San Diego: Academic Press.

Berntson, G. M. and J. D. Aber. 2000. Fast nitrateimmobilization in N-saturated temperate forestsoils. Soil Biology and Biochemistry 32: 151–156.

Burgi, M., E. W. B. Russell, and G. Motzkin. 2000.Effects of postsettlement human activities on forest composition in the northeastern UnitedStates: a comparative approach. J. Biogeography27:1123–1138.

Boose, E. R., K. E. Chamberlin, and D. R. Foster.2001. Landscape and regional impacts of hurri-canes in New England. Ecological Monographs 71:27–48.

Catovsky, S. 2000. Linking community dynamicsand ecosystem function in mixed conifer broad-leaved forests. Ph.D. Thesis, Harvard University.

Cavender Bares, J. and F. A. Bazzaz. 2000. Changesin drought response strategies with ontogeny inQuercus rubra: implications for scaling fromseedlings to mature trees. Oecologia 124: 8–18.

Compton, J. E. and R. Boone. 2000. Long-termimpacts of agriculture on organic matter poolsand nitrogen transformations in central NewEngland forests. Ecology 81: 2314–2330.

Eberhardt, R. 2001. Implications of land-use legaciesin the sand plain vegetation of Cape Cod,National Seashore. M.F.S. Thesis, HarvardUniversity.

Foster, D. R. 2001. Conservation lessons and chal-lenges from ecological history. Forest History Today,

Fall 2000: 2–11.Foster, D. R. 2000. Using history to interpret current

environmental conditions and future trends: anexample from the U.S. Long Term EcologicalResearch (LTER) Program. PAGES Newsletter 8:23–25.

Foster, D. R. 2000. Linking the deep and recent pastto the modern New England landscape. Rhodora102: 278–279.

Foster, D. R. 2001. New England’s Forest Primeval.Wild Earth 11: 40–44.

Francis, D. R. and D. R. Foster. 2001. Response ofsmall New England ponds to historic land use.The Holocene 11: 301–312.

Francis, D. R. 2001. Reconstructing past hypolim-netic oxygen conditions in a temperate multide-pression lake using chemical evidence and chi-ronomid remains. J. Paleolimnology 25: 351–365.

Golub, A. 2000. Russian forests for climate changemitigation: an economic analysis. Environmentand Natural Resources Program, J. F. K. Schoolof Government, Harvard University, Cambridge,MA.

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Hall, B. R., D. J. Raynal, and D. J. Leopold. 2001.Environmental influences on plant species com-position in ground-water seeps in the CatskillMountains of New York. Wetlands 21: 125–134.

Jenkins, J. C., D. W. Kicklighter, and J. D. Aber.2000. Predicting the regional impacts ofincreased CO

2and climate change on forest pro-

ductivity: a model comparison using PnET-IIand TEM 4.0. Pp. 383–423 in R. Birdsey, J.Hom, and R. Mickler (Eds.), Responses of NorthernForests to Environmental Change. Springer-Verlag.

Johnson, T. C., S. L. Barry, Y. Chan, and P.Wilkinson. 2001. Decadal record of climate vari-ability spanning the past 700 years in theSouthern Tropics of East Africa. Geology 29:83–86.

Kicklighter, D. W., M. Bruno, S. Dönges, G. Esser,M. Heimann, F. Helfrich, F. Ift, F. Joos, J. Kaduk,G. H. Kohlmaier, A. D. McGuire, J. M. Melillo,R. Meyer, B. I. N. A. Moore, I. C. Prentice, W.Sauf, A. Schloss, S. Sitch, U. Wittenberg, and G.Würth. 2000. A first-order analysis of the poten-tial role of CO

2fertilization to affect the global

carbon budget: a comparison study of four ter-restrial biosphere models. Tellus 51B: 343–366.

Kohyama, T., E. Suzuki, T. Partomihardjo, and T.Yamada. 2001. Dynamic steady state of patch-mosaic tree-size structure of a mixed dipterocarpforest regulated by local crowding. EcologicalResearch 16: 85–98.

Mack, R. N., D. Simberloff, W. M. Lonsdale, H.Evans, M. Clout, and F. A. Bazzaz. 2000. Bioticinvasions: causes, epidemiology, global conse-quences and control. Ecological Applications 10:689–710.

Magill, A. H. and J. D. Aber. 2000. Dissolved organ-ic carbon and nitrogen relationships in forest lit-ter as affected by nitrogen deposition. Soil Biologyand Biochemistry 32: 603–613.

Magill, A. H. and J. D. Aber. 2000. Variation in soilnet mineralization rates with dissolved organiccarbon additions. Soil Biology and Biochemistry 32:597–601.

Magill, A. H., J. D. Aber, G. M. Berntson, W. H.McDowell, K. J. Nadelhoffer, J. M. Melillo, andP. A. Steudler. 2000. Long-term nitrogen addi-tions and nitrogen saturation in two temperateforests. Ecosystems 3: 238–253.

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Martin, M. E. and J. D. Aber. 2000. Estimating forestcanopy characteristics as inputs for models of for-est carbon exchange by high spectral resolutionremote sensing. Pp. 61–72 in H. Gholz (Ed.), TheUse of Remote Sensing in the Modeling of ForestProductivity at Scales from the Stand to the Globe. TheNetherlands: Kluwer Academic Publishers.

Minocha, R., S. Long, A. H. Magill, J. Aber, and W.H. McDowell. 2000. Foliar free polyamine andinorganic ion content in relation to soil and soilsolution chemistry in two fertilized stands at theHarvard Forest. Plant and Soil 222: 119–137.

Motzkin, G. 2001. Montague Plain: a resource man-ual of historical and ecological information gath-ered by Harvard Forest. Report to the EcologicalRestoration Program, MA Biodiversity Initiative,MA Natural Heritage and Endangered SpeciesProgram.

Orwig, D. A., C. V. Cogbill, D. R. Foster, and J. F.O’Keefe. 2001. Variations in old-growth struc-ture and definitions: forest dynamics onWachusett Mountain, Massachusetts. EcologicalApplications 1: 437–452.

Parshall, T. and R. Calcote. 2001. Effect of pollenfrom regional vegetation on stand scale forestreconstruction. The Holocene 11: 81–87.

Parshall, T. 2001. Book Review: R. Judd (1997)Common Lands, Common People: The Originsof Conservation in Northern New England.Ecology 82: 1497–1498.

Tomlinson, P. B. 2001. Biology of Trees Native to TropicalFlorida, second edition. Privately printed.

Tomlinson, P. B. and U. Posluszny. 2001. Genericlimits in the seagrass family Zosteraceae. Taxon50: 429–437.

Tomlinson, P. B. and E. H. Zacharias, 2001.Phyllotaxis, phenology and architecture inCephalotaxus,Torreya and Amentotaxus (Coni-ferales). Botanical Journal of the Linnean Society 135:215–228.

Tomlinson, P. B., J. B. Fisher, R. E. Spangler, and R.A. Richer. 2001. Stem vascular architecture in therattan palm Calamus (Arecaceae-Calamoideae-Calaminae). American Journal of Botany 88:797–809.

Yano, Y., W. H. McDowell, and J. D. Aber. 2000.Biodegradable dissolved organic carbon in forestsoil solution and effects of chronic nitrogen dep-osition Soil Biology and Biochemistry 32: 1743–1751.

ACKNOWLEDGMENT OF SUPPORT

Activities described in this Annual Report are supported by funds provided by the followingsources, to which we are indebted:

Friends of the Harvard ForestCommonwealth of Massachusetts

Department of Environmental Management

Division of Fisheries and WildlifeBiodiversity InitiativeEcological Restoration programNatural Heritage and

Endangered Species ProgramAndrew W. Mellon FoundationNational Science Foundation

Biological Field Stations and Marine

LaboratoriesEcosystem StudiesInternational ProgramsLong-term StudiesEcology ProgramResearch Experience for

UndergraduatesThe Family of Richard Thornton FisherU.S.D.A. Competitive Research Grants,

ForestryDepartment of Energy — NIGECSouthern Taconics Research and

Conservation Center

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GIFTS

The Library at Harvard Forest welcomed more giftsof books to its collection. Brian Hall, a ResearchAssistant, donated eight books, which included LucyBraun’s Deciduous Forests of Eastern North America. SarahCooper-Ellis, a former staff member, donated bookson Plant Hormones, Plant Development, VegetationDynamics, and Global Change. Jim Baird has againdonated later issues of The Auk.

Aimee Peterson donated to the archive map col-lection a copy of the Petersham Historical Society’s“Historical Map of Petersham,” which was originallycreated with the help of the Harvard Forest.

NEW FUNDING

D. R. Foster. Long Term Ecological Research atHarvard Forest (LTER III). National ScienceFoundation. $4,255,000.

D. R. Foster, D. A. Orwig, and D. B. Kittredge.Forest Response to the Introduced Hemlock WoollyAdelgid: Development of Silvicultural Guidelines.U.S.D.A. Northeastern Area Federal Focus Funding.$35,000.

D. R. Foster and J. Hadley. Evaluating the effects ofdiverse vegetation types and soil drainage classes onnet carbon exchange of a landscape mosaic withmobile and fixed eddy covariance systems. NIGEC.$165,000.

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David R. Foster Petersham, MassachusettsDirector August 2001

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THE HARVARD FOREST 2000–2001Harvard University