THE HARVARD FOREST 2002-2003

Harvard University

Front Cover: Sunset from the Prospect Hill FireTower

Back Cover: Leap O’ Faith

Photography by David Foster, Jakara Hubbard, Ava Foster, John O’Keefe, and Betsy Colburn

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ANNUAL REPORT OF THE HARVARD FOREST 2002-2003

Contents

Personnel at the Harvard Forest - - - - - - 4Introduction to the Harvard Forest - - - - - - 5New Staff - - - - - - - - 6Research Activities - - - - - - - - 7Bullard Fellows - - - - - - - - 27Educational Activities - - - - - - - - 30

Summer Research Program - - - - - - 30Activities of the Fisher Museum - - - - - - 32

Meetings, Conferences, Seminars - - - - - 33Forest Management and Maintenance - - - - - - 34Library and Archives - - - - - - - - 35Information Management and Technology Advancement - - - 36Activities of the Harvard Forest Staff - - - - - - 36Visiting Research Scientists - - - - - - - 40Publications - - - - - - - - - 41Acknowledgment of Support and Gifts - - - - - - 45New Funding - - - - - - - - - 45

http://harvardforest.fas.harvard.edu

Audrey Barker Plotkin Research AssistantLeann Barnes Laboratory TechnicianSylvia Barry Musielewicz Research AssistantEmery Boose Information and Computer

System ManagerJeannette Bowlen AccountantJohn Burk Archivist and Librarian Philip Burton Bullard FellowJessica Butler Research AssistantRichard Cobb Research AssistantElizabeth Colburn Bullard FellowWillard Cole Woods CrewAnthony D’Amato Graduate StudentElaine Doughty Research AssistantAshley Eaton LandscaperEdythe Ellin Director of

AdministrationAaron Ellison Senior EcologistAdrian Fabos Facilities ManagerEd Faison Research AssistantSamantha Farrell Laboratory TechnicianBarbara Flye Librarian and SecretaryRichard Forman Landscape EcologistCharles H. W. Foster AssociateDavid Foster DirectorPeter Franks Bullard FellowKelli Graves Secretarial AssistantLucas Griffith Woods CrewJulian Hadley EcophysiologistBrian Hall Research AssistantJulie Hall Research Assistant Linda Hampson Staff Assistant

Alice Ingerson Bullard FellowSultana Jefts Research AssistantHolly Jensen-Herrin Research AssistantMatthew Kelty Bullard FellowDavid Kittredge Forest Policy AnalystPaul Kuzeja Research AssistantOscar Lacwasan Woods CrewDavid Lindenmeyer Bullard FellowHeidi Lux Research AssistantDana MacDonald Research AssistantGlenn Motzkin Plant EcologistJohn O’Keefe Museum and Schoolyard

CoordinatorDavid Orwig Forest EcologistJulie Pallant System and Web

AdministratorSarah Parnes Graduate StudentLaura Pustell Research AssistantFrancis Putz Bullard FellowDorothy Recos Smith Staff AssistantMichael Scott Woods CrewJudy Shaw Woods CrewNavjot Sodhi Bullard FellowRachel Spicer Graduate StudentBernhard Stadler Bullard FellowKristina Stinson Research AssociateP. Barry Tomlinson E. C. Jeffrey Professor

of Biology, EmeritusBetsy Von Holle Post-doctoral FellowJohn Wisnewski Woods CrewSteven Wofsy Associate

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PERSONNEL AT THE HARVARD FOREST 2002-2003

INTRODUCTION TO THE HARVARD FOREST

Since its establishment in 1907 the Harvard Foresthas served as Harvard Univerisity’s rural laboratoryand classroom for research and education in forestbiology and ecology. Through the years researchershave focused on forest management, soils and thedevelopment of forest site concepts, the biology oftemperate and tropical trees, plant ecology, forest eco-nomics, landscape history, conservation biology, andecosystem dynamics. Today, this legacy of activities iscontinued as faculty, staff, and students seek tounderstand historical and modern changes in theforests of New England and beyond resulting fromhuman and natural disturbance processes, and toapply this information to the conservation, manage-ment, and appreciation of natural ecosystems. Thisactivity is epitomized by the Harvard Forest LongTerm Ecological Research (HF LTER) program,which was established in 1988 through funding bythe National Science Foundation (NSF).

Physically, the Harvard Forest is comprised ofapproximately 3,000 acres of land in the north-central Massachusetts town of Petersham that includemixed hardwood and conifer forests, ponds, streams,extensive spruce and maple swamps, fields, anddiverse plantations. Additional land holdings includethe 25-acre Pisgah Forest in southwestern NewHampshire (located in the 5000-acre Pisgah StatePark), a virgin forest of white pine and hemlock thatwas 300 years old when it blew down in the 1938Hurricane; the 100-acre Matthews Plantation inHamilton, Massachusetts, which is largely comprisedof plantations and upland forest; and the 90-acre TallTimbers Forest in Royalston, Massachusetts. InPetersham a complex of buildings that includesShaler Hall, the Fisher Museum, and the John G.Torrey Laboratories provide office and experimentalspace, computer and greenhouse facilities, and lec-ture room for seminars and conferences. Nine addi-tional houses provide accommodations for staff, vis-iting researchers, and students. Extensive records,including long-term datasets, historical information,original field notes, maps, photographic collections,and electronic data are maintained in the HarvardForest Archives.

Administratively, the Harvard Forest is a depart-ment of the Faculty of Arts and Sciences (FAS) ofHarvard University. The Harvard Forest administersthe Graduate Program in Forestry that awards a

masters degree in Forest Science and faculty at theForest offer courses through the Department ofOrganismic and Evolutionary Biology (OEB), theKennedy School of Government (KSG), and theFreshman Seminar Program. Close association is alsomaintained with the Department of Earth andPlanetary Sciences (EPS), the School of PublicHealth (SPH), the Graduate School of Design (GSD)at Harvard, and the Department of Natural ResourceConservation at the University of Massachusetts, theEcosystems Center of the Marine Biological Labora-tory, and the Complex Systems Research Center atthe University of New Hampshire.

The staff and visiting faculty of approximatelyfifty work collaboratively to achieve the research,educational, and management objectives of the Har-vard Forest. A management group meets monthly todiscuss 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 an infu-sion of outside perspectives. The six-member WoodsCrew and Facilities Manager undertake forest man-agement and physical plant activities. The Coordi-nator of the Fisher Museum oversees many educa-tional and outreach programs.

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

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NEW STAFF

Betsy Von Holle began a three-year post-doctoralposition working with David Foster, and GlennMotzkin supported by a National Parks EcologicalResearch fellowship. Her research focuses on the linkbetween disturbance history and habitat invasibilityby exotic plants at Cape Cod National Seashore.Betsy received her B.S. from the University ofCalifornia at San Diego and her Ph.D. from theUniversity of Tennessee.

Jess Butler and Dan Atwater

Five new research assistants began work at theForest. Jessica Butler, who recently received an MSfrom Oregon State University and has a B.S. fromthe University of Montana, will be conductingresearch on pitcher plant ecosystems with AaronEllison. Edward Faison who recently completed aM.S. in Botany from UVM and has a B.A. fromConnecticut College, will be documenting the long-term history of hemlock in New England with DavidFoster.

Sultana Jefts and Laura Pustell will be conductingresearch on the hemlock woolly adelgid with DaveOrwig. Sultana has an MS from the University ofMaine and B.S. from Evergreen State College. Laurahas a B.S. from the University of Connecticut. PaulKuzeja, who is conducting research on forest carbonexchange with Julian Hadley, received a M.S. fromthe University of Vermont, a M.Ed. from UMass-Amherst, and a B.A. from Amherst College.

Tony D’Amato

Tony D’Amato, a Ph.D. student in the Forestryprogram at University of Massachusetts as anadvisee of Dave Orwig, is conducting research on thedynamics of old-growth forests in westernMassachusetts. Tony received his M.S. at OregonState University and a B.S. from the University ofMaine. Richard Shulhoff, currently Deputy Directorof the Arnold Arboretum at Harvard University, wasaccepted into the M.F.S. program working withDavid Foster and Dave Orwig.

Club Paleo: Dana MacDonald, Ed Faison, Elaine Doughty, Sylvia Barry Musielewicz

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RESEARCH ACTIVITIES

Community Assembly in a Changing World

With new support from NSF, Aaron Ellison isexpanding his research on wetland communityassembly, disassembly, and reassembly followinglarge-scale disturbance. One project focuses on thestructure of invertebrate communities inhabiting thenorthern pitcher plant, Sarracenia purpurea (see photo), acarnivorous plant that grows in bogs from Florida toCanada. The plant’s pitcher-shaped leaves fill withrainwater into which ants, flies, moths, and spidersfall and drown. Unlike its congeners, S. purpurearelies on a small aquatic food web consisting of bac-teria, protozoa, rotifers, mites, and larvae of mosqui-toes, midges, and flesh flies to digest the arthropod

prey that it captures and to mineralize the nitrogenlocked up in the prey. In the last several decades, asconcentrations of nitrogen in rainfall have increased,pitcher plants in New England have come to rely lesson carnivory as their needs for nitrogen are met moreand more by nitrogen in rainfall (see graph). Currently,Aaron, Jess Butler, and REU student Dan Atwaterare investigating experimentally how much nitrogenplants obtain from prey verus how much they obtainfrom precipitation. This project is aimed at develop-ing a complete nitrogen budget for S. purpurea, andunderstanding the consequences of this shift for theassembly and dynamics of the aquatic food web inthe face of long term, persistent environmentalchange.

Pitcher plants in the western United States face adifferent challenge. In the Siskiyou Mountains ofsouthern Oregon, the endemic Darlingtonia californicais restricted to serpentine fens. During the summer of2002, the Biscuit Fire, one of the largest wildfires in

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Aaron Ellicson

Response of the northern pitcher-plant, Sarracenia purpurea, todirect nutrient additions. Shape changes are expressed as twocorrelated characters: relative tube diameter (an indication ofhow large an opening the leaf has in which it can capture prey)and relative keel size (an indication of the relative size of thepitcher tube itself, in which prey are digested). The jointchanges in shape are illustrated with line drawings of a normalpitcher (no nutrients added) with a relatively large mouth andsmall keel (top left) and a flattened leaf (phyllode) with a rela-tively small mouth and large keel (bottom right). Joint means arelabeled with the nutrient addition treatment. The plot at thetop illustrates the N and P concentrations added for each ofthe nine treatments (N added in gray; P in black). Theseresults illustrate that leaf shape is sensitive to added N, but notto added P. Figure from Ellison & Gotelli (2002) PNAS 99:4409–4412.

Pitcher Plant

the history of the Pacific Northwest, burned nearly500,000 acres in the center of the Darlingtonia’s rangein southwestern Oregon and northern California.Building on three years of pre-fire data, Aaron andhis colleagues Nick Sanders (Humboldt State), andErik Jules (Humboldt State) received funding fromNSF to begin a study of the re-assembly of these fencommunities.

Population and Community Studies of InvasivePlant Species in New England

A group including Kristina Stinson and KathleenDonohue from OEB and Betsy Von Holle from theHarvard Forest are investigating population, commu-nity, and landscape-level interactions between nativeand invasive plants in Massachusetts. The majorobjectives are to: identify links between land use andthe presence/abundance of non-natives; develop pre-dictive models of invasion based on historical andpresent-day levels of anthropogenic and natural dis-turbance; test the direct impact of invasive plantcover on the survival and performance of nativeflora; and identify evolutionary potential for furtherspread within invading populations. Major results to date have focused on the biennial herbaceousplant, garlic mustard (Alliaria petiolata) (see photo). ThisEurasian exotic is a persistent invader in NewEngland forests where it threatens native understoryand tree seedling species. We have documented neg-ative relationships between A. petiolata and importantnative tree seedlings, such as sugar maple and ash,in western Massachusetts.

Current research is focused on competitionbetween native species and two important exoticinvaders, garlic mustard and barberry (Berberis thun-bergii), using experiments in which the exotic speciesare fully or partially removed from forest habitats.We have also begun conducting demographic sur-veys and evolutionary studies of garlic mustard pop-ulations in forested, intermediate, and sunny habitattypes in newly invading populations at HarvardForest. Our preliminary data suggest that seedlingnumbers are much higher in sunny habitat, but thattransition rates between seedling, adult, and repro-ductive stages are higher in intermediate and foresthabitat than in open habitat. However, plants insunny habitat are much larger, have higher photo-synthetic rates and seed productivity than plants inthe other two habitats. These results indicate thatgarlic mustard populations in high-light habitats aremore likely to spread than those in low-light habitats,an important consideration for population dynamicsin forests with different levels of canopy openness.New reciprocal transplant studies at Harvard Forestare underway that will address the role of maternalhabitat on the germination, development, and fitnessof offspring in garlic mustard in different sites.Combined with long-term, statewide data on land-use history and habitat characteristics, this researchwill enable us to understand how evolutionary andpopulation dynamics may contribute to invasionfrom population-to-landscape scales.

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Flower of Garlic Mustard (Alliaria petiolata)

Student Gui Woolston counts seedlings, rosettes andreproductive individuals in a plot

This summer, EunSuk Kim, a graduate studentin OEB, and Kathleen Donohue set up an experi-ment in the new experimental garden plot with thehelp of many of the REU summer students. Thisexperiment will measure the influence of the seasonof seed maturation on the germination timing andlife-history expression of five weeds. These weeds areknown to exhibit variation in life-history schedules inways that may influence their generation time andconsequent population growth rates. The studies areaimed at identifying the contributions of maternalenvironmental effects, germination timing, and life-history expression to weedy behavior.

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Alex Sanchez-Sierra, Christian Foster, Luke Durbin

Kathleen Donohue and Deberat Perez-Rivera

The planted gardenPrepping the site

Vegetation Dynamics of Ridgetop Pitch Pine and Red Pine Communities

Dave Orwig

Dave Orwig, Glenn Motzkin, and David Foster areconducting a study of the vegetation and long-termdynamics of ridgetop sites in southern New Englandthat support uncommon pitch pine or red pine com-munities. The object of this study is to understandthe long-term history and dynamics of these unusualhabitats and vegetation types in order to guide theirprotection and conservation into the future. This pastyear, we focused our efforts on several summits in thesouthern Taconic Mountains with dwarf pitch pinevegetation, as well as sites in western Massachusettswith native red pine. In our previous work on Mt.Everett in southwestern Massachusetts, we foundthat the dwarf pitch pines ranged up to ~ 170 yearsold, although the average age was substantiallyyounger. The pitch pine “trees” on these summits arefrequently only 1–3 m (3–10 feet) tall, and oftenexhibit highly contorted shapes. Occasionally thetrees are completely prostrate. The importance ofhardwoods, the proportion of dead versus live pitchpine, and evidence of past fire varies greatly across

the sites. Interestingly, although we could find noevidence of fire in the historical record or soils on Mt.Everett, a fire that occurred east of the summit insummer 2002 smoldered for approximately sevenweeks before finally being extinguished by rains. Todate, all ridgetops that we’ve visited in the regionwith native red pine have evidence of past fire, andseveral sites have extensive recent red pine moralityfrom undetermined causes.

Patterns of Forest Harvesting in Massachusetts

Timber harvesting is the most widespread andimportant form of disturbance to New Englandforests and yet relatively little is known about the fre-quency, type, spatial distribution, or ecological con-sequences of this activity. Assessing forest harvestingas an ecological process is especially difficult in thisregion due to the large proportion of private landownership, the small size and selective nature ofmany logging operations, and the general inability toidentify harvested areas through satellite imagery orother forms of remote sensing. In order to evaluatethe ecological and conservation implications of mod-ern forest cutting activities David Foster, DaveKittredge, and Glenn Motzkin have initiated astatewide research project across Massachusetts.With the cooperation of staff from the MassachusettsDepartment of Environmental Management (DEM)and assistance from Sweetwater Trust, John Burkgathered data this year on harvesting patterns acrossthe Commonwealth since 1984 using regulatory datacollected on all commercial logging operations byDEM.

Glenn Motzkin

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In collaboration with Frank Lowenstein, AndyFinton, and Mark Anderson from The NatureConservancy, we will use these data to: (1) documentthe temporal and spatial variation in forest harvestingacross the state; (2) analyze the distribution of log-ging with regard to important conservation valuesand physical, cultural, and environmental factors; (3)evaluate harvesting impacts on critical ecologicalcharacteristics, including invasive species distribu-tion, native plant species richness and composition,and soil characteristics; and (4) compare harvestingpatterns and intensity with natural disturbanceregimes across the region. Methodologies and resultsfrom this study will advance conservation planningacross the eastern U.S., and will have broad rele-vance to global forest policy and conservation effortsaddressing such issues as carbon dynamics and theinvasion of exotic species. For instance, in a pilotstudy of the North Quabbin region, Dave Kittredge,Andrew Finley, and David Foster found a surprising-ly high frequency and unexpected pattern of cutting:approximately 1.5% of the forest area was loggedannually in a spatially random pattern that has leftfew large forest blocks undisturbed over the pasttwenty years.

Forest Policy

Dave Kittredge finished his sabbatical year awayfrom the University of Massachusetts by completinghis study of private forest owner cooperation in coun-tries with developed economies and temperateforests. He identified examples of successful coopera-tion in over nineteen countries, involving millions of

owners and hectares. Many examples of cooperationhave been ongoing for decades. These were compiledin a report to the Ford Foundation’s Community-Based Forestry Project, and have been condensedinto a manuscript for journal publication. Addi-tionally, Dave wrote an article on his 2002 visit toSweden, where he studied formal, private, forestowner cooperatives.

In the summer of 2002, Kittredge and summerstudent Katy Nicholson began a study of the patternsof timber harvesting throughout Massachusetts.Based on the statewide compilation of timber harvestregulatory data, this study seeks to identify the loca-tions where harvesting ceases to occur in otherwiseforested communities. What combination of factors(e.g., population density, amount of forest, level ofaffluence, road density, parcel size) leads to theabsence of harvesting as an activity in communities,that are otherwise as much as 40–50% forested? Thestudy is being expanded in the summer of 2003through the efforts of REU student Joanna Bate, toinclude data from New Hampshire.

Temperate Forest Interactions with theAtmosphere

The forest and other ecosystems exchange gases andother materials with the atmosphere; understandingthe relative amounts of material entering and leavingterrestrial ecosystems is a critical aspect of the envi-ronmental research undertaken at the HarvardForest. A major part of these investigations is under-

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Dave Kittredge

C. H. W. Foster

taken at the Harvard Forest EnvironmentalMeasurement Station (HFEMS), which consists ofan instrumented tower and an array of ecologicalplots. Trace gas concentrations and fluxes betweenthe atmosphere and the forest are measured at thetower. At the plot sites we measure biomass incre-ment, collect foliar and litter samples, and trackchanges in coarse woody debris (CWD).

2002 marked the beginning of the second decadeof carbon (CO2) flux measurements. Although theNet Ecosystem Exchange (NEE) of carbon hasremained fairly steady at 2.2 ton C ha-1 y-1, the GrossEcosystem Exchange (GEE), or uptake, and Eco-system Respiration (R) have been generally increas-ing as the forest biomass increases with time.

During 2002 we began a new study with col-league Neil Pedersen at Lamont Doherty Obser-vatory to compare tree-ring cores from the plots withdiameter increment and eddy flux measurements.

As a part of her Senior Thesis at HarvardCollege, Wendy Liu, measured the flux of CO2 fromdowned CWD with direction from PostdoctoralFellow David Bryant. She found that total CO2 fluxfrom CWD was nearly equal to soil respiration.Residence time of carbon in the CWD pool was esti-mated at 8.3 to 10.5 years for the Simes Tract. Inputsof CWD from normal tree mortality and minor dis-turbance such as windthrow, lightning, ice damage,and insects are currently being measured and willprovide additional information on turnover rates forthis important portion of the carbon budget.

Results from investigations by REU studentGreg Santoni suggest that soil respiration follows adaily trend, responding to soil temperature and mois-ture. This temporal variation in CO2 flux and spatialvariation in the driving variables prompted collabo-ration with Patrick Crill and Ruth Varner of theUniversity of New Hampshire. Eight automatedCO2 measurement chambers were installed in spring2003 at the margin of the Bigelow swamp northwestof the HFEMS tower. Soil moisture and temperatureare also being measured every two hours at eachchamber.

David Bryant is managing the network ofautochambers and is taking manual respiration meas-urements weekly within the swamp. The combineddataset will allow landscape level analysis of the spa-tial and temporal pattern of soil respiration in wet-land, upland, disturbed and undisturbed forests.

Air quality measurements are a major focus ofstudies at the HFEMS. An automated gas chromato-graph, one of several similar instruments in a U.S.National Oceanic and Atmospheric Administration(NOAA) global network, is used to track changes inthe concentration of chlorofluorocarbons (CFC) nowthat their production and use has been stopped.Although the concentration of this important green-house gas is trending downward as expected, ourobservations suggest that emissions are still occur-ring, presumably from residues in old equipment,landfills, and stockpiled material.

Measurements of ozone and other air pollutantshave been made at HFEMS since 1990. During thesummer of 2002 we observed an unusual event withvery high carbon monoxide (CO) concentrationsalong with thick haze from a group of forest fires innorthern Quebec in early July. Peak concentrationsof CO approached 800 ppb, which greatly exceedsthe typical summertime values. In August the Har-vard Forest experienced a multi-day episode of highozone.

Variation in Carbon Exchange Among Forests

During the past year Julian Hadley and Paul Kuzejaused an eddy flux system similar to that at theHFEMS to complete the first full year of carbonexchange measurements on Little Prospect Hill(LPH), in a red oak and red maple forest that devel-oped after an intense fire in 1957. This site is higher,drier, and supports a much younger forest than thearea around the HFEMS tower site. Measurements

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Zachary Liscow, Kathryn McKain, Lucy Huytra, andSteve Wofsy

at both sites (as well as measurements from the old-growth hemlock stand reported on in last year’sannual report) are designed to assess the variation incarbon exchange among different forest types as itrelates to forest characteristics as well as climatic vari-ation and change. These issues are important as NewEngland’s forests affect global carbon exchange,atmospheric CO2 concentrations, and the green-house effect resulting from globally rising atmos-pheric CO2.

For the months of May through December 2002for which we have analyzed the data, net carbonexchange followed a similar pattern to the older andlower oak-maple forest near the HFEMS tower, buttotal carbon storage was lower on Little ProspectHill. However, the pattern of carbon storage differsstrikingly between the hardwood forests and the old-growth hemlock forest. Whereas carbon storage in

the hemlock stand fell sharply between May andAugust, it rose during that period to its highest levelsof the year at the HFEMS. Considering the monthsof May through October, the period when deciduoustrees have photosynthetically active foliage, the twodeciduous forest sites showed a positive relationshipbetween monthly average soil temperature and car-bon storage. In contrast, in the old-growth hemlockforest, monthly carbon storage declined with increas-ing soil temperature. Because monthly average soiltemperature is closely linked to monthly average airtemperature (although less variable), the results sug-gest that carbon storage in oak-maple forests mayactually be enhanced by warmer summers. At least,carbon storage in oak-maple forests is unlikely todecline sharply in warm summers, as will probablyoccur in old hemlock forests.

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Chris Graham and Julian Hadley using aScholander pressure chamber to measure leaf

water potentials after a carbon dioxideexchange measurement

Preparing a black birch leaf forwater potential measurement

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Net monthly carbon exchange of two different forests measured by eddy covariance, compared to the 60–100-year-old forest around the Harvard Forest Environmental Measurement Site

(ftp://ftp.as.harvard.edu/pub/nigec/HU_Wofsy/hf_data.) Left: an old eastern hemlock forest; Right: a younger, deciduous forest on a dry site that developed after a forest fire in 1957.

Both deciduous forest sites are dominated by red oak and red maple.

Left: Relationship between monthly average soil temperature and monthly net carbon exchange during April through October in the hemlock forest and the 60–100-year-old deciduous forest around the HFEMS.

Right: The same relationship for the two deciduous forests during May through October 2002.

Analytical Web – Collaboration betweenEcologists and Computer Scientists

In order to develop the values for carbon exchangefrom the eddy flux towers, researchers combinemany thousands of measurements and estimatesthrough a complex series of computations. Docu-menting such analyses in order to preserve themethodology and to provide a means of replicatingthe procedure is a major, though generally under-appreciated, challenge in ecological science. In 2002Harvard Forest scientists joined with colleagues inthe Department of Computer Sciences at theUniversity of Massachusetts to develop tools for cap-turing and facilitating sequences of complex datatransformations. Working with new computer soft-ware developed at UMass we have begun to recordthe models and data through a process and productreferred to as the “Analytic Web.” Using the AnalyticWeb, other scientists will be able to precisely replicatethe process of model development, and to quicklyand easily test the effects of changes in input data orin the structure of models that convert raw data intohigher-level information such as regional forest car-bon exchange estimates. Carbon exchange modelingis but one of numerous potential uses of the AnalyticWeb, since most scientific studies involve sequencesof data manipulation and analysis. The NationalScience Foundation’s Information TechnologyResearch (ITR) program is supporting the AnalyticWeb project, including data collection and carbonexchange modeling work at the Harvard Forest, andcomputer software development at the University ofMassachusetts.

Soil Warming Experiments

During the past year we activated our latest warmingexperiment in the Slab City Tract of the HarvardForest — The Barre Woods Megaplot. This long-termexperiment funded by the National Institute forGlobal Environmental Change (NIGEC) and LTERis designed to analyze some of the consequences ofglobal climate change on the soil environment. Aftertwelve years of warming the soil in the Prospect Hilltract, we have expanded the scope of our experimentfrom a replicated set of 6 x 6m plots to a pair of 30 x30m plots — one heated and one control. At this newsite, we are exploring the possible fates of increasedcarbon and nitrogen moving through the northernhardwood forest system and potentially into the

atmosphere. The past decade plus of warming onProspect Hill suggests that both the carbon and nitro-gen cycles are stimulated by warming, but that theseeffects are transient. With our new experiment, wehope to answer the question: Does the forest makeuse of the additional nitrogen mobilized in the systemand store more carbon as a result?

Heidi Lux

Mapping and experimental installation tookplace in 2001, and a year of baseline measurementsof soil nutrient dynamics and tree growth began inthe spring of 2002. Heating began in May of 2003and sampling continues. Early soil respiration meas-urements reveal similar increases due to heating towhat we observed on Prospect Hill.

Jerry Melillo and Paul Steudler presented prelim-inary data from the new experiment as well as resultsfrom our twelve-year study at meetings of the Eco-logical Society of America and the AmericanGeophysical Union in 2002, and at the annual Har-vard Forest Symposium in 2003. In December of2002, we published a paper in Science, “Soil Warmingand Carbon-Cycle Feedbacks to the Climate System”(Melillo et al., 2002), presenting a synthesis of ourwork at the Prospect Hill Soil Warming Site. In thesummer of 2002, REU student Nicole Nowinski,from Carleton College, worked with us on a nitrogenbudget for the new site. Margaret Graham, fromDartmouth College, is working with the group forthe summer of 2003.

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Long-term Climate Change – Signals for theLittle Ice Age in New England?

The “Little Ice Age,” ca. A.D. 1450 to 1850, was anapparently global climatic period characterized in theNorthern Hemisphere by highly variable conditions,frequent long winters, and short, cool summers. Innorthwestern Europe environmental signals of thisperiod are recorded in glacial activity, coastal sedi-ments, tree rings, and historical documents, whichindicate a period of cool and variable weather includ-ing shorter growing seasons and increased stormi-ness. More recently, glacial, tree ring, lake level, and

historical data indicate coincident changes in terres-trial and aquatic ecosystems in North America.

The low-resolution and poor temporal control ofmost pollen and stratigraphic data has restrictedreconstructions of subtle paleoclimatic changes inNew England. Furthermore, the end of the “Little IceAge” coincides with the period of European settle-ment and broad-scale deforestation, which lead tomajor changes in pollen abundances. However, somestudies suggest that significant environmental andvegetation changes occurred prior to settlement. Forexample, our recent studies of the southern NewEngland landscape determined that the major

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Outflow from Harvard Pond

changes since European settlement, including thedecline in beech and hemlock and the regionalhomogenization of vegetation composition, wereactually initiated some 300–500 years beforeEuropean arrival.

In the current study, we combine pollen datafrom many new sites and the Harvard Forest archiveswith detailed historical land-use and land cover mapsto analyze vegetation and environmental change inMassachusetts over the past 2,000 years. Massa-chusetts is a particularly good study region because itwas colonized early, is covered by unique historicalrecords, and embraces a wide range of vegetation,environments, and cultural conditions. We analyzedrecords from over twenty ponds arrayed across thestate’s gradients in physiography, climate, geology,and natural disturbance. Chronological control wasprovided through lead-210 and radiocarbon datingand sediments were sampled at high-resolution inter-vals. Small (<10 ha) and primarily spring-fed kettlelakes were selected to emphasize the local to sub-regional vegetation signal. Historical data includearchaeological information and pre-settlement vege-tation derived from witness trees records in earlyland surveys.

Preliminary results suggest that the effects of the“Little Ice Age” varied geographically, with the mag-nitude and nature of vegetation changes differing byvegetation type and ecoregion. A decrease in beechand hemlock is most evident in high-elevation sites,while low-elevation sites of oak-hardwood forestcomposition show relatively little change during thepast 1,000 years. In many cases, changes in forestcomposition dynamics are accompanied by coinci-dent changes in aquatic vegetation and percentorganic matter, presumably reflecting changes inwater balance and lake levels.

Integrated Hemlock Studies

At the Harvard Forest, we are anticipating witnessingthe disassembly and reassembly of our hemlockforests as the hemlock woolly adelgid continues itsrelentless northward march. This year, we found theadelgid for the first time in the hemlock stand onProspect Hill as well as on scattered trees aroundShaler Hall. A major research thrust focused on thebiological, hydrological, and physical changes associ-ated with the loss of hemlock and its replacement bybirch has begun with support from NSF (to DaveOrwig and David Foster) and from the Harvard

University Center for the Environment (to a largegroup of scientists from the Department ofOrganismic and Evolutionary Biology, Earth andPlanetary Sciences, Kennedy School of Government,and the Harvard Forest).

Various studies led by Dave Orwig are investi-gating the various impacts of the introduced hemlockwoolly adelgid on New England forests. This effortexpanded with new NSF funding in 2003 supportingtwo new research assistants, Sultana Jefts and LauraPustell. This summer Dave, Sultana, and summerstudent David Franklin continued to sample perma-nent plots established at eight sites in Connecticutduring 1997 to examine the effect of hemlock declineand mortality on the timing and extent of changes inthe nitrogen cycle. After several years of HWA infes-tation, tree crowns continue to deteriorate, allowingmore light to reach the forest floor. In addition,infested stands have lower surface soil moisture lev-els, higher soil temperatures, and higher net nitrogenmineralization and net nitrification rates than unin-fested stands or stands with low damage. Resin bags

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Prospect Hill hemlock forest

buried in the soil at damaged sites also capturedhigher amounts of ammonium and nitrate than atuninfested stands, indicating that nitrogen is becom-ing more available as a result of the microenviron-mental changes associated with HWA damage. Wewill continue to sample these stands to examine thelong-term changes in N cycling associated with hem-lock deterioration and eventual replacement by hard-wood species such as black birch and oak. Local highschool student Leann Barnes, assisted with many ofthe laboratory activities associated with these labor-intensive studies.

Dave, Sultana, and Audrey Barker Plotkin con-tinued to collect baseline information on soil nitrogencycling and microenvironments in the Prospect Hilllong-term hemlock grid in anticipation of the arrivalof HWA. The many past research projects and therich data that exist for this site provide an unusualopportunity to document the current and future forest changes with the detailed knowledge of pastforest dynamics.

Over the last four years we have been interestedin changes in decomposition associated with thedecline of hemlock due to HWA, because these maybe closely linked to observed changes in nitrogencycling. Under the direction of Richard Cobb, wehave studied three potential drivers of decompositionassociated with HWA arrival and the decline of hem-lock: (1) changes in the chemistry in leaf litter; (2)changes in microclimate; and (3) hemlock replace-ment by black birch. Foliar carbon and lignin did notappear to be affected by HWA infestation, but foliarnitrogen (% N) was higher in infested stands. In turn,this increased the rate of N immobilization in decom-posing foliage, likely leading to the increased N avail-ability mentioned above. Microclimatic changes,involving increased light and temperature led tomore rapid sub-surface decomposition and initiallyslower decomposition on the forest floor. Finally,results after six months show that black birch litterdecomposed more rapidly than hemlock or mixed lit-ter. We will continue this study for another year to

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Soils in infested hemlock sites in Connecticut typically had higher net n-mineralization rates than control stands (a),and as stands continued to deteriorate, higher net nitrification rates (b).

Resin bags incubated in soils captured higher amounts of ammonium and nitrate in infested versus control hemlock stands, indicating greater N availability.

examine longer-term dynamics as these substratesbreak down over time.

A new project was initiated by Dave, LauraPustell, and summer student Amanda Park to exam-ine the vegetation and ecosystem consequences ofhemlock removal by logging, the primary manage-ment response to the HWA. Permanent plots arebeing established in logged hemlock sites inMassachusetts to compare the vegetation, microenvi-ronment, and nitrogen cycling responses to thosetriggered by HWA infestations.

As a expansion of a large-scale project examiningHWA distribution in Connecticut, Dave Orwig andsummer students Nick Povak and Don Niebyl havebegun sampling hemlock stands across Massa-chusetts. We have now mapped the distribution ofhemlock stands in a broad transect acrossConnecticut and Massachusetts (see map). During thesummers of 2002–3, we sampled over seventy ofthese stands in Massachusetts and collected informa-tion on forest structure and composition, crownvigor, site characteristics, presence of HWA, and theextent and spatial patterns of damage generated bythis insect since its arrival in 1989. Almost 50% ofstands sampled had HWA, although overstory hem-lock mortality remained low. This information willbe incorporated into a GIS analysis of landscape-level, biological, edaphic, and historical factors thatcontrol the damage patterns observed in hemlock.Summer student Joe Brown, under the direction Dr.Scott Costa from the University of Vermont, visited

many of the Massachusetts sites and conducteddetailed sampling of hemlock branches to provideaccurate estimates of HWA densities and to helpdevelop a protocol for sampling HWA populations.

Bernhard Stadler initiated an experiment withDavid Orwig and Richard Cobb to study the effectsof trophic interactions between HWA and epiphyticbacteria on the spatial and temporal variability inenergy and nutrient flow through the hemlockcanopy at sites varying in HWA infestation level.Uninfested hemlock showed a significant decline incanopy foliar biomass from the center of a tree to theperiphery. In contrast infested trees, had significantlyless canopy biomass, exhibited no trend in canopybiomass, and produced less new foliage. Infestedtrees had significantly higher foliar % N, with N high-est in young foliage where HWA was at the highestdensity. Epiphytic microorganisms showed little dif-ference in abundance on needles growing in differentparts of the canopy, but bacteria, yeasts, and filamen-tous fungi thrived on medium and heavily infestedcompared to uninfested trees. The amount of raincollected in throughfall varied spatially beneath unin-

19

Transect through southern New England for the study ofhemlock woolly adelgid. The extent of hemlock forest

increases dramatically in Massachusetts.

Dead hemlock trees on Mt. Tom

fested trees, with most throughfall passing in theperiphery of the canopy and least close to the trunk.No gradient occurred beneath infested trees, throughwhich relatively more precipitation was percolating.Throughfall chemistry was strongly affected byHWA infestation, as higher concentrations of differ-ent nitrogenous compounds and carbon as well asdifferent ion species were leaching from infestedcanopies to the forest floor. We intend to continuethese investigations in the future to help identify andunderstand the chain of mechanisms that lead tochanges in decomposition and nutrient availability asforests respond to invasive pests.

In a related effort last summer, student JackieGuzman and Audrey Barker Plotkin studied seedlingbank dynamics of eastern hemlock at two contrastingsites in the Forest. The study sought to document theage structure of the hemlock seedling bank, comparegrowth rates between sites, and determine whetherrecruitment into the seedling bank is episodic or con-tinuous. Seventy seedlings less than 4.5 feet in heightwere collected in both an old hemlock forest and a50-year-old mixed hardwood forest. Height, basaldiameter, and age were determined for each seedling.Average ages differed between sites: thirty years oldin the hemlock stand and fourteen years old in thehardwood forest. The oldest individual was forty-sixyears old. Both sites displayed an exponential growthrate, suggesting that seedlings slowly and continu-ously grow into larger size classes. Analysis of age

structure indicates that recruitment into the seedlingbank is continual but episodic. These results suggestthat hemlocks use a seedling bank as a regenerationstrategy and that further study of hemlock seedlingbank dynamics might aid future restoration projects.

Long-term, Large-scale Hemlock Experiment

To complement Dave’s regional studies David Foster,Aaron Ellison, and Audrey Barker Plotkin haveestablished a set of eight, 90 x 90m experimentalplots in the Simes Tract in southern Petersham. Six ofthese plots are dominated by hemlock; two plots willbe logged (a typical treatment for many hemlockstands across the region); all the hemlocks in anoth-er two plots will be girdled (to simulate some aspectsof adelgid infestation, including generation of coarsewoody debris); and the remaining two plots willserve as controls. Two additional plots already sup-port mature hardwood forests and thereby representpost-adelgid “controls.” We plan to collect pre-treat-ment (and pre-adelgid) baseline data for two sum-mers before we log or girdle the plots. During thesummer of 2003, Eric Davidson and KathleenSavage from the Woods Hole Research Center beganto measure soil respiration in the plots; Dave Orwigand his team began studies of litter decompositionand nutrient fluxes; and Aaron Ellison and two REUstudents, Jonathan Chen and Matt Lau, inventoriedthe diversity and structure of ant communities in thehemlock and hardwood stands.

Hydrological Studies

We received funding from NSF and HarvardUniversity to begin long-term hydrological measure-

20

Audrey Barker Plotkin

Alana Belcon

ments on two small headwater streams in theProspect Hill Tract. The first stream drains the BlackGum Swamp and adjacent woodlands and flows nearShaler Hall on its way to Nelson Brook, the MillersRiver, and ultimately the Connecticut River. The sec-ond stream drains the steep valley to the west ofProspect Hill and flows through the wetland north ofthe HFEMS Tower on its way to Bigelow Brook, theSwift River, the Chicopee River, and the ConnecticutRiver. Separating the two streams in the middle of theProspect Hill Tract is the watershed boundarybetween the Millers River and Chicopee Riverbasins.

Permanent weirs are planned for each stream,whose watersheds, though adjacent and comparablein size, differ significantly in topography, soils,hydrology, land-use history, stream biota, and forestvegetation (including the abundance of hemlock).

Each station will be provided with electricity and acommunications link to Shaler Hall. Heating cablesand hoods will be used to prevent ice buildup at theinstrumented sites in winter. Measurements, includ-ing surface discharge and water temperature, will beposted on the Harvard Forest Web page in near-realtime and submitted monthly to the LTER NetworkHydroDB database. Piezometers and soil moisturesensors will be deployed above the weirs to measuresoil moisture and subsurface water flow.

Botanical Studies

Comparative studies of the biology of palms haveemphasized the distinctiveness of those with a climbing habit, most familiar in the rattans, thesource of canes used widely in the furniture industry.Continuing with this research, an earlier analysis byMartin Zimmermann was resurrected, completed,and published, dealing with the South Americangenus Desmoncus, a rattan analogue. It demonstratesthe considerable structural constraints on axial andappendicular water supply. This research is the struc-tural compliment to extensive field research by grad-uate student Alex Cobb, working in Malaysia andQueensland.

Having demonstrated unusual cytological prop-erties of tension fibers in the tropical genus Gnetum,further research is aimed at establishing if these prop-erties extend to the tension wood fibers of dicotyle-dons. These cells demonstrate remarkable dynamicaspects of anatomy in woody plants that are of con-cern to the timber industry. The cells develop ten-sions that can re-erect bent stems in trees.

Additional research, in part using the collections

21

Debarat Perez-Rivera

Field work in Tom Swamp

of the Arnold Arboretum, has centered on the dis-tinctive cone morphology of the “taxads,” a group ofconifers, most familiarly represented by Taxus, all ofwhich have fleshy animal dispersed seeds. Clearlythe reproductive anatomy of these conifers representsconsiderable reduction so that the presumed ances-tral coniferous cone is difficult to discern. The genusTorreya, which includes several threatened species, inparticular has been very incompletely analyzed. Areview of conifers overall demonstrates the diversityof reproductive mechanisms in a lineage often over-looked because of their constant wind-pollination.

Other research with Dr. Usher Posluszny at theUniversity of Guelph, Ontario, Canada has demon-strated first, that flowers of the New Caledoniangenus Amborella, considered to be the basal-most line-age of flowering plants has, in fact, a floral morphol-ogy that can scarcely be seen as “ancestral” to otherflowering plants. Secondly, attention has been drawnto establishment growth in seedlings of seagrasses(marine angiosperms) as an important biologicalphase that has remained unexplored by morpholo-gists and ecologists. How do the seeds of such plantsbecome established in shallow but tidal environ-ments, amidst shifting sediments? The axis in oneexample (Zostera) seems to act like a kedge anchor.

Xylem Physiology of Forest Trees

N. Michele Holbrook of the Department ofOrganismic and Evolutionary Biology (OEB) andmembers of her lab are continuing studies of thestructure and function of trees, with an emphasis onwater flow through the xylem. Cavitation sets ahydrodynamic limit to water transport and thus is acritical process affecting tree function. Studies under-way this year address the link between xylem struc-ture and vulnerability to cavitation, from the level ofsingle bordered pits to the hydraulic architecture ofwhole trees. A project led by Maciej Zwieniecki andBrendan Choat focuses on the biophysics of cavita-tion and repair. This work involves detailed studiesof water flow through stems in response to increasingxylem tension. Summer REU students Nora Lahrand Chris Petit are conducting independent projectsrelated to xylem function. Nora’s project investigateshow the cavitation thresholds of individual xylemvessels vary as a function of age and position withinlarge sugar maple trees; meanwhile Chris is compar-ing water flow through healthy and adelgid-infectedhemlock branches to test the hypothesis that hemlock

wooly adelgid infection reduces the water transportcapacity of the xylem. A second major line of workfocuses on the ability of changes in xylem sap ionconcentrations to alter xylem hydraulic resistancethrough their effect on the pectin hydrogels in pitmembranes. Studies ongoing this year are designedto test the hypothesis that recirculation of K+ ionsfrom phloem to xylem may allow plants to dynami-cally affect their hydraulic architecture, providingplants with a mechanism to enhance water move-ment to the most productive branches.

Rachel Spicer, a Ph.D. student in OEB, has beenstudying the process of sapwood senescence thatleads to heartwood formation in forest trees. A majorquestion in sapwood physiology is how the livingcells within woody stems are supplied with oxygen.The respiration of these cells is an important deter-minant of total stem respiration. In order to testwhether internal stem conditions are ever limiting tothese cells, Rachel has been measuring oxygen con-centrations at different radial depths within stems.She is also measuring rates of sapwood respirationunder different O2 and CO2 environments, andcomparing this response across different ages of sap-wood. Results suggest that heartwood formation isunlikely to be driven by anoxia, or oxygen depriva-tion, both because oxygen contents in the oldest sap-wood rarely get that low, and because respirationremains unaffected by decreasing oxygen down toabout 2%. Summer REU student Teresa Abbott isworking with Rachel to determine how CO2 con-centrations influence sapwood respiration.

A NSF and U.S. Depatment of Agriculture

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Brady Hardiman using the canopy lift (aka Bucky) to take leaf measurements

(USDA) supported workshop on “Long DistanceTransport Processes in Plants” was held at theHarvard Forest during October 2002. Organized byMissy Holbrook and Maciej Zwieniecki, this threeday work shop was attended by over seventy partici-pants from more than ten countries. Proceedingsfrom the workshop will be published by AcademicPress as part of their Physiological Ecology series.

Harvard Forest Tract Maps

Since 1908, researchers at Harvard Forest havemapped and conducted inventories of the forests onits three largest tracts approximately every 10 to 20years. Features such as soils, elevation, damage fromthe 1938 hurricane, and silvicultural treatments weremapped. Over the past several months, Brian andJulie Hall have been involved in digitizing these mapsin a standardized and readily usable format. Thesedata will be useful in the selection and description of study sites and will be important in documentingthe influence of past forest management activities and environmental conditions on Harvard Forestproperties.

23

Oxygen contents within stems at different radial depths, measured in spring and summer.

The effect of high CO2, which is known to be elevatedwithin stems, on parenchyma cell respiration in whitepine (Pinus strobus) sapwood. Inner sapwood is about

ten years older than outer sapwood.

Brian Hall

1830 Map Series

We have finished digitizing and compiling the 1830map series, which documents the landcover and cul-tural features in Massachusetts near the peak of agri-cultural land use. Features shown include forests,roads, and major buildings (including forest-relatedtypes such as sawmills and tanneries). These datahave been widely distributed to research and conser-vation groups and have been incorporated into thestate-wide conservation plans by the Commonwealthof Massachusetts.

Both the 1830 and Harvard Forest Tract datalay-ers will be available for viewing and downloadingfrom the Harvard Forest Web site.

Harvard Forest LTER Program

The Harvard Forest is one of twenty-four 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-wide

activities 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, K. Donohue), Earth andPlanetary Sciences (S. Wofsy, B. Munger), andHarvard Forest (D. R. Foster, D. Kittredge, G.Motzkin, D. Orwig, A. Ellison, B. Colburn, E. Boose,J. Pallant) at Harvard University as well as theMarine Biolbogical Laboratory (MBL)-EcosystemsCenter ( J. Melillo, P. Steudler), University of NewHampshire ( J. Aber, A. Magill, S. Ollinger. S. Frey),University of Massachusetts (M. Mulholland, E.Chilton), and Brandeis University (B. Donahue).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-termand intensive measurements of forest and plant struc-ture, function, and dynamics; (3) experimentalmanipulations; and (4) synthesis and modeling. TheHarvard Forest Ecology Symposium is held to pres-ent current research of the LTER and NationalInstitute for Global Environmental Change(NIGEC) programs, with abstracts published annu-

24

Julie Hall

John Aber provides an overview to the LTER visiting committee

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

National Institute for Global EnvironmentalChange

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 and

observational 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 inaddition to faculty from the State University of NewYork (D. Fitzjarrald), Woods Hole Research Center(E. Davidson), University of California (S.Trumbore), U.S. Geological Survey (E. Sundquist),and Harvard Forest ( J. Hadley).

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David Foster outlines research to the LTER visiting committee

HARVARD FOREST ECOLOGY SYMPOSIUM 2003 (*denotes summer student)

J. Aitkenhead-Peterson et al. Dissolved Organic Carbon and Nitrogen in a Hardwood Forest Floor.M. Albani and P. Moorcroft. Modeling the Impact of Hemlock Loss on New England Forests.D. Barnes et al. Greenhouse and Ozone-Depleting Gases in Rural New England. S. Barry, D. R. Foster and B. Hall. The Little Ice Age in New England.W. Borken et al. Experimental Drought Effects on Soil Respiration in a Temperate Forest Soil.P. J. Burton, C. M. Burton et al. Vegetation in a Forest 20 Years After Creation of Small Forest Gaps.R. Cobb et al. Hemlock Woolly Adelgid Effects on Foliage and Decomposition.E. A. Colburn. Vernal Pools in the New England Landscape.E. A. Colburn and H. Jensen-Herrin. Monitoring Freshwater Macroinvertebrates on Cape Cod.E. A. Colburn and D. A. Orwig. Effects of Hemlock Decline on Forest Stream Communities.E. A. Colburn, J. Choiniere et al. Headwater Habitat Streams in Central Massachusetts.B. Dail et al. Results from a Wet NH4NO3 Canopy Fertilization in a Maine Spruce-Hemlock Forest.K. Donohue, S. Takao* et al. QTL Analysis of Parental Effects on Dormancy in Arabidopsis thaliana.K. Donohue, N. Wender* et al. Genetic Architecture and Dispersal Plasticity in Arabidopsis thaliana. A. M. Ellison and N. J. Gotelli. Effects of Nutrient Stress on a Co-evolved Food Web.A. M. Ellison et al.. Biology of Loss and Replacement of a Core Species.P. Franks, M. Zwieniecki, and N. M. Holbrook. The Regulation of Transpiration Rate in Plants.S. D. Frey and M. Knorr. Nitrogen and the Structure and Function of the Soil Microbial Community.J. Gaudinski et al. The Importance of Belowground Plant Allocation for Soil Carbon Sequestration.J. Hadley. Carbon Exchange in a 55-Year-Old Oak-Maple Forest on a Dry Upland Site in 2002.B. Hall, G. Motzkin, D. R. Foster, M. Syfert, and J. Burk. 300 Years of Forest Change in Massachusetts. L. Hutyra, Santoni*, S. Pears* et al. Carbon Cycling at the Harvard Forest, a Bottom-up Approach.A. E. Ingerson. Using History for Land Management in New England.D. B. Kittredge, D. R. Foster, J. Burk and G. Motzkin. Logging as Disturbance in Massachusetts.D. Köste, et al. Paleolimnological Assessment of Human-Induced Impacts on Walden Pond.X. Lee, H. Wu, J. Sigler, and C. Oishi. Rapid and Transient Response of Soil Respiration to Rain.W. Liu et al. Respiration of Coarse Woody Debris in Central Hardwood Stands.P. J. Melcher et al. Vulnerability of Xylem Vessels to Cavitation in Acer saccharum (Marsh).J. M Melillo, H. Lux et al. Soil Warming: The Fist Decade and the Megaplot Experiment.G. Motzkin, D. A. Orwig and D. R. Foster. Vegetation and Disturbance in Ridgetop Pine Communities.G. Motzkin et al. Conservation of Uncommon Plant Communities: the Northeastern Coastal Region.J. W. Munger, J. Budney, and S. C. Wofsy. Air Quality at the Harvard Forest: Pollution Events in 2002.K. L. Musgrove* and E. A. Colburn. Distribution of Malacostracan Crustaceans in Vernal Pools. J. O’Keefe. Woody Species Phenology, Prospect Hill Tract, Harvard Forest — 2002.S. V. Ollinger and J. D. Aber. Effects of Multiple Environmental Changes on Forest Carbon Exchange.D. Orwig and D. R. Foster. Landscape-Level Analyses of Hemlock Woolly Adelgid Outbreaks.D. Orwig, R. Cobb, M. Kizlinski, and D.R. Foster. Ecosystem effects of Hemlock Woolly Adelgid.S. Parnes. Distribution and Abundance of hemlock and white pine since 1733 in Petersham.N. Pederson et al. Relation between Eddy-Flux Measurements and Tree-Ring Estimates of Growth. J. G. Quijano and A. P. Barros. Incorporating Vegetation Dynamics into a Hydrological Model.L. Sack, P. D. Cowan, N. Jaikumar, and N. M. Holbrook. The ‘Hydrology’ of Leaves.J. Sigler et al. Emission and Long-Range Transport of Mercury from a Boreal Fire.T. W. Sipe. Responses by Herbs and Seedlings to Enriched Carbon Dioxide and Irradiance Regimes.R. Spicer. Heartwood Formation and Patterns of Cell Death in Woody Stems.B. Stadler et al. Adelgid impact on Throughfall Chemistry and Microorganism Abundance.K. A. Stinson. Does Selection on Ecophysiological Traits Facilitate Invasion of Alliaria petiolata?S. Urbanski et al. Carbon Sequestration at Harvard Forest in 2001. A Banner Year?B. Von Holle et al. Disturbance Histories as a Predictor of Habitat Invasibility in a Mosaic Landscape.

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BULLARD FELLOWS

Philip Burton (Symbios Research, Smithers, B.C.,Canada), with assistance from his wife, Carla, resam-pled fifteen experimental canopy gaps (30 m2 to 720m2) on the Prospect Hill Tract. This silviculturalexperiment was initiated in 1982 by David Hibbs,and had been sampled previously by Tim Sipe in1984. Twenty years after logging most gaps weredominated by red maple and red oak stump sproutsand by black birch seedlings. Tree populations hadundergone considerable self-thinning from the densi-ties encountered two growing seasons after logging.Many trees greater than 10 m tall were found in thelarger gaps, while the understory vegetation was nowlargely similar to that in the undisturbed forest.

Phil completed editorial work on a book entitledTowards Sustainable Management of the Boreal Forest.Forthcoming from NRC Research Press in Sep-tember 2003, the volume summarizes the first sevenyears of research conducted under the auspices ofCanada’s Sustainable Forest Management Network.He also completed a paper on a theory of forest standedge effects, and book chapters on landscape ecolo-gy, conservation biology, and forest restoration inBritish Columbia. Phil was an active participant inthe Harvard Forest Lab Group and Fakhri Bazzaz’slab discussions in Cambridge. During one of thesnowiest winters in recent years, the Canadians oftenbroke trail for cross-country skiing enthusiasts at theForest.

Betsy Colburn (Harvard Forest) analyzed habitatcharacteristics of Cape Cod wetlands with regards topopulations of a rare, northern caddisfly, Phanoceliacanadensis, a species associated with Sphagnum-domi-nated sites. Betsy sampled Sphagnum wetlands else-where in Massachusetts in search of other popula-tions of this caddisfly, worked with Emery Boose onplanning for installation of permanent weirs on twoHarvard Forest streams, initiated research on head-water habitat streams in central Massachusetts, andwith colleagues from Wachusett Meadow WildlifeSanctuary, Mount Wachusett State Reservation, andAntioch/New England Graduate School, mappedeighteen intermittent headwater streams, assessed in-stream habitat, and started biological sampling forsalamanders and aquatic invertebrates. She analyzedvernal pool samples from across the state with twosummer students in 2002. REU student KateMusgrove presented a poster at the North AmericanBenthological Society meeting, examining crustacean

distributions in vernal pools. Betsy initiated studies of streams in hemlock- and

hardwood-dominated watersheds as baseline forlong-term evaluation of the effects of hemlock declineon stream ecology. She participated in discussionswith several agencies and interested private partieson how restoration of tidal flow into PilgrimLake/East Harbor, Truro, might affect nuisancemidge outbreaks from the lake. A long-term moni-toring protocol for freshwater macroinvertebrates inlakes, ponds, and vernal pool wetlands in the CapeCod National Seashore was initiated. With HollyJensen-Herrin, research assistant, literature reviewswere carried out; gradients of chemistry, physicalvariables, limnological status, and historic land usewere evaluated; and preliminary field sampling toevaluate sampling methods and identify appropriatebiological indices for long-term monitoring was con-ducted, and analysis of samples initiated.

Betsy conducted public workshops on vernalpools (Cape Cod); intermittent stream hydrology,biology, and sampling (central Massachusetts); andthe connections between groundwater, vernal pools,freshwater wetlands, and kettle ponds (Cape Cod).She presented a paper on “History as a habitat vari-able for aquatic systems” in plenary session of NewEngland Association of Environmental Biologists’

27

Phil and Carla Burton

Annual Meeting in Westminster, Mass., and servedon the Masters Thesis committee for SalvatoreBeatini, Worcester Polytechnic Institute. She also participated in U.S. EPA’s New England BiologicalAssessment of Wetlands Work Group (NEBA-WWG), contributed to deliberations of NorthAmerican Benthological Society’s Science and PolicyCommittee and Conservation Committee and con-tributed to planning for Massachusetts NaturalHeritage and Endangered Species Program’s AquaticBiomap Project. Betsy focused on revising a bookmanuscript on the ecology and conservation of ver-nal pools in northeastern North America, which hasbeen accepted for publication, as well as one peerreviewed manuscript.

Peter Franks ( James Cook University) carriedout experiments to determine the role of leafhydraulic conductance in the stomatal control of leafgas exchange. In collaboration with Missy Holbrookand Maciej Zwieniecki, new techniques utilizing thecell pressure probe were employed to obtain data onthe hydraulic conductance between leaf veins andstomatal guard cells. These data have improved themechanistic modeling of stomatal function. Peterpublished four research articles and co-authored abook chapter on stomatal control of water loss fromtrees. He was an invited speaker at the FifthInternational Workshop on Field techniques inEnvironmental Physiology (Organized by theUniversity of Edinburgh, U.K. and held in Tenerife,Spain, March 2003), where he presented two keynotetalks on new techniques for studying stomatal func-tion and water flow in woody plants.

Alice Ingerson (Waban, Mass.) conducted overforty-five interviews around New England, to deter-mine whether and how conservationists and landmanagers were using historical ecology to make andevaluate practical decisions. The interviews suggest-ed that historical ecology is currently used fairly nar-rowly: to set targets for ecological restoration (usual-ly, the state of the landscape prior to European set-tlement); and in interpretive programs for the publicfocused on landscape features as cultural or historicresources (most often cellar holes, stone walls, andabandoned roads). Yet most interviewees themselvesalso questioned these applications, recognizing thatpre-European ecological conditions were not neces-sarily more stable or preferable to conditions in otherperiods; and that the entire New England landscape,including the distribution of vegetation and wildlifeas well as stonewalls and cellar holes, is a culturalartifact. When discussing these criticisms, manyinterviewees endorsed very different potential uses ofhistory: as a set of methods for analyzing present andfuture as well as past landscape change, and as a setof skills that, if taught more widely, could foster moreeffective public participation in long-term, adaptivemanagement. Alice will organize a researcher/practi-tioner roundtable at Harvard Forest in the fall of2003, to explore recommendations and implementa-tion based on these findings.

Alice organized a mini-symposium, “Making ItReal: Bridging the Gap Between Historic andWorking Farm Landscapes in New England,” andspoke on “ ‘Farm’ Is a Verb: Interpreting Change inNew England’s Working Landscapes,” Ninth AnnualDeerfield-Wellesley Conference on American Culture(Historic Deerfield, Massachusetts). She spoke andled discussions on “A Critical User’s Guide toChange and Stability in Cultures and Ecosystems,”for students in the University of Massachusetts atAmherst’s Department of Landscape Architecture,and for staff of the Trustees of Reservations and theMassachusetts Audubon Society; assisted the NewEngland Small Farm Institute and the University ofMassachusetts at Boston with grant proposals onapplied farm history and urban environmental citi-zenship; organized a session, “Research as Activism:Participatory Approaches to EnvironmentalHistory;” and served as chair and discussant for“Nature Conservation and Historic Preservation:The Landscape as Connection,” American Societyfor Environmental History (Providence, RhodeIsland). Alice developed and taught the new course,

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Alice Ingerson

“Place, Community, and Time: Social andEnvironmental History for Landscape Practitioners,”Landscape Design Program of the ArnoldArboretum, Harvard University, and spoke as part ofa panel on “The Future of Local History,” Bay StateHistorical League (Lowell, Massachusetts).

Matthew Kelty (University of Massachusetts atAmherst) spent much of his time working on a newedition of the text The Practice of Silviculture, which heis coauthoring with Mark Ashton and Bruce Larson.One of his main efforts in this tenth edition of thebook has been to incorporate the idea of using natu-ral forest dynamics as a model for designing silvicul-tural systems. He also worked on a number ofresearch papers, one of which dealt with the use ofsludge from municipal wastewater treatment as fertil-izer for red pine. The results of that study support thefindings of the Harvard Forest chronic nitrogen dep-osition project, in that red pine showed reduced

growth rates in response to nitrogen fertilization inboth cases. Matt began a field project with JohnO’Keefe and UMass student Helen Johnson dealingwith the long-term management plots in Tom SwampII, which had been established in 1907; this includesdesigning a thinning that is consistent with the pastmanagement practices. Matt presented a seminar atthe Harvard Forest “Ecological restoration of forestsheavily browsed by deer: the problem of hay-scentedfern,” and led a two-week field trip for forestry stu-dents from Germany and the U.S. to the NorthernRocky Mountain region.

David Lindenmayer (The Australian NationalUniversity) completed several papers on the impactsof salvage logging on post-disturbance stand recoveryand biodiversity response (Australian Forestry). He alsoreviewed the value of natural disturbance-basedapproaches as guidelines for forest management.Other papers completed included a review of biodi-versity conservation in plantation systems (BiologicalConservation), analyses of landscape supplementationby birds in an Australian plantation landscape(Conservation Biology), reptile response to grazing andplantation establishment (Journal of Applied Ecology),the value of nestboxes for cavity-dependent fauna(Wildlife Research), population recovery of small mam-mals following a population removal experiment(Journal of Applied Ecology), the use of systematicreviews in medical sciences as a model for conserva-tion application (Biodiversity and Conservation), and anew landscape model for tracking biodiversityresponse to landscape change (Oikos). He finished co-editing a book on the impacts of fire in Australianecosystems (CSIRO Publishing) and well advancedthe major update of a textbook on PracticalConservation Biology (Surrey Beatty and Sons). Hepresented talks at the University of California, Davis,the Wildlife Conservation Society (New York),Weyerhauser Company (Vancouver), University ofMaine, and the Yellowstone to Yukon Conference(Calgary).

Bullard Fellows for 2003–2004 include Drs. IanBaillie (Bedford, England), Paul Barten (Universityof Massachusetts), Julia Jones (Oregon StateUniversity), John Klironomos (University of Guelph,Ontario, Canada), Antonia Lara (UniversidadAustral de Chile), Matts Lindbladh (SwedishUniversity of Agricultural Science), and ThomasSinclair (University of Florida).

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David Lindenmayer

EDUCATIONAL ACTIVITIES

David Foster was joined by John O’Keefe, GlennMotzkin, and Dave Orwig in leading the HarvardForest Freshman Seminar this spring. The seminarmeets over four weekends to provide an overview ofHarvard Forest research and an introduction to thelandscape and history of New England. It finisheswith the students undertaking an independent study.David also taught two sections of Biology 299 ForestPractice and Research, a seminar for advancedundergraduates and graduate students that was orig-inally developed by Ernie Gould.

Sarah Parnes is continuing her master’s researchon the history and dynamics of hemlock and whitepine in southern New England. She is joined in theMFS program by Richard Shulhoff, Deputy Directorof the Arnold Arboretum. Richard is beginning a

study that examines options for the management ofHemlock Hill at the Arnold Arboretum, a distinctiveand historical hemlock grove that is heavily infestedby the hemlock woolly adelgid.

For the past two years Barry Tomlinson has beeninvolved in an unusual course, sponsored by theKenan Foundation, that seeks to facilitate knowledgeabout tropical plants and how such information maybe incorporated into the teaching of introductorybiology courses, the participants coming from col-leges and universities throughout the United Statesand Canada. The course is taught on Kauai, Hawaii,in July, with a follow-up in Miami in February.

Summer Research Program

The Harvard Forest Summer Student Research pro-gram, coordinated by Edythe Ellin and assisted byLaurie Miskimins and Jakara Hubbard, attracted adiverse group of students to receive training in scien-tific investigations, and experience in long-term eco-logical research. All students worked closely withresearchers while many conducted their own inde-pendent studies. The program included weekly sem-inars from resident and visiting scientists, discussionson career issues in science, and field exercises onsoils, land-use history, and plant identification. Anannual field trip was made to the Institute ofEcosystem Studies (Millbrook, N.Y.) to participate ina Forum on Careers in Ecology. Students presentedmajor results of their work at the Annual SummerStudent Research Symposium in mid-August.

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John O’Keefe

Freshman Seminar Students Deberat Perez-Rivera, Alana Belcon, Zachary Liscow

Summer Students 2003

Teresa Abbott Willamette UniversityDaniel Atwater University of KansasJoanna Bate Haverford CollegeAlana Belcom Mount Holyoke CollegeJoseph Brown Holyoke Community CollegeJonathan Chen Oberlin CollegeNaomi Clark W. Virginia UniversityJennifer Clowers Franklin & Marshall CollegeLuke Durbin Illinois Wesleyan UniversityDavid Franklin Lehigh UniversityChristopher Graham Dickinson CollegeMargaret Graham Dartmouth CollegeBrady Hardiman Ashland UniversityKatherine Joseph Virginia Tech UniversityEleanor Lahr Ithaca CollegeErin Largay Yale UniversityMatthew Lau Humboldt State UniversityZachary Liscow Harvard UniversityRebecca Lohnes Yale UniversityNick Malizia Clark University

Kathryn McKain Mount Holyoke CollegeKatie Musgrove SUNY, UlsterJulia Nelson Stanford UniversityDonald Niebyl Virginia Tech UniversityNicole Nowinski Carleton CollegeAmanda Park SUNY, SyracuseDaberat Perez-Rivera Metropolitan University,

Puerto RicoChristopher Petit Carleton CollegeNick Povak Virginia Tech UniversityAlex Sanchez-Sierra Franklin & Marshall CollegeJulie Vuong Franklin & Marshall CollegeKristin Wilson University of Maine, OronoWilliam Woolston Harvard University

Assistant Program CoordinatorsLaurie MiskiminsJakara Hubbard

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ACTIVITIES OF THE FISHER MUSEUM

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

Just in time for summer visitors in 2003 our newinterpretive signs for the Natural History Trailthrough Jonathan Sanderson’s Farm were installed.These striking aluminum signs, designed in collabo-

ration with Michael Connors of the Inform Group inNorthampton, combine line graphics and photoswith text to enable visitors to interpret the history ofthe land as well as more recent management andresearch activity. The interpretation on the signsserves as an excellent extension of the lessons fromthe dioramas onto the landscape itself.

In November the Museum volunteers celebratedthe completion of another very successful weekendschedule at our twelfth Volunteer RecognitionDinner, at which Mary Ann Walker once againreceived special thanks for her continuing, enthusias-tic work as volunteer coordinator. A familiar group,including Walt Davidson, Martha Siccardi, Bill andMarianna Berry, Roger and Barbara Corey, BobLane, and Dick Sherwood received special recogni-

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Tim Sipe, Alex Sanchez-Sierra, Jen Clowers, and Julie Vuong

Jakara Hubbard and Laurie Miskimins

Matthew Lau and Jonathan Chen

Chris Petit

tion for being the most active volunteers during theseason. The group was deeply saddened in the fall bythe death of Bob Reed, a long-time friend and volun-teer, with an immense fondness for Harvard Forest,and again in the spring by the loss of friend and vol-unteer Ed Wierman.

During the year the Museum provided programsfor twenty-six elementary and secondary schools,

thirty-six college and university classes, and fourteencommunity and professional groups. In July theForest hosted the forestry training for the nationalCanon Envirothon. This high school student compe-tition, hosted by Hampshire College in 2002,brought more than fifty student/teacher teams, repre-senting almost all the states and many Canadianprovinces, together for a week of education andexamination in a range of environmental disciplines.

Meetings, Conferences, Seminars

In October the Forest hosted an international confer-ence on Long Distance Transport in Plants, organ-ized by Missy Holbrook of the Department of Organ-ismic and Evolutionary Biology. The FourteenthAnnual Harvard Forest Long-Term EcologicalResearch Symposium and National Institute forGlobal Environmental Change meeting was held inthe Museum on February 12. Other meetings at theHarvard Forest included meetings of the Massa-chusetts Extension Service Coverts Project,Massachusetts Department of EnvironmentalManagement Logging Workshop, MassachusettsForest Stewardship Committee, Massachusetts StateForestry Committee, Massachusetts ForestryAssociation, New England Forestry Foundation,Mount Wachusett Community College, North-eastern Forest Soils Conference, North QuabbinRegional Landscape Partnership, and VegetationControl Service. The Forest also hosted a facultyretreat for the Harvard University Department ofOrganismic and Evolutionary Biology.

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Michael Connors, Adrian Fabos, and John O’Keefefollowing the installation of new trail signs

Speakers in the Harvard Forest Seminar series included:

Robert Askins Connecticut CollegeBarbara Bedford Cornell UniversityDavid Bryant Harvard UniversityCarla Burton Symbios Research and

RestorationPhil Burton Symbios Research and

Restoration John Confer Ithaca CollegePeter Franks James Cook University

(Queensland, Australia) Peter Frumhoff Global Resources and

Environment Program, Union of Concerned Scientists

Lauren Howard University of NewHampshire

Yongsong Huang Brown UniversityMatthew Kelty University of

MassachusettsThom Kyker-Snowman Mass. Metropolitan

District CommissionAlice Ingerson Arnold Arboretum Walt Landgraf Stone Museum at People’s

State Forest, Barkhamsted, Conn.

James Levitt Program for ConservationInnovation, Harvard Forest

David Lindenmayer Australian NationalUniversity

Michelle Manion Global EnvironmentProgram, Union of Concerned Scientists

Leslie Mehrhoff George Safford TorreyHerbarium, University of Connecticut

Christian Messier Université de Québec àMontréal

Paul Nickerson U.S. Fish and WildlifeService

Keith Nislow USDA Forest Service, Northeastern Research Station

Graeme Patterson Wildlife ConservationSociety

Jan Rowan U.S. Fish and WildlifeService

Scott Shumway Wheaton CollegeBill Sobczak Holy Cross CollegeDennis Souto USDA Forest ServiceBernhard Stadler University of Bayreuth Tibor Standovar Eoetvoes Lorand

University, HungaryChloe Stuart Mass. Natural Heritage

and Endangered SpeciesProgram

P. B. Tomlinson Harvard ForestBetsy Von Holle National Park Ecological

Research Fellow, Harvard Forest

Claudia Zimmermann Université Laval, Québec

FOREST MANAGEMENT AND MAINTENANCE

Adrian Fabos and his talented staff, better known asthe “Woods Crew,” supported the research efforts ofthe Harvard Forest community, which included up-

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Lucas Griffith, Mike Scott, Woody Cole,and John Wisnewski

grading the two walk-up scaffold towers (providingaccess to the forest canopy and above); training andsupporting a multitude of staff and visitingresearchers in the safe operation of Harvard Forest’snewly acquired mobile canopy lift — capable of reach-ing up seventy feet; building a boardwalk into abeaver pond to support research on gas exchange;

and converting a field below the Lyford House into acultivated research garden for botanical studies and experimental research. The greenhouse atTorrey Lab was made operable after many years ofinactivity and currently supports several research initiatives.

There were many other support efforts to theeducational and research programs during the pastyear, including improvements to facilities, labs andhousing. Lyford House, used for Bullard Fellows andother researcher housing needs, was completely de-leaded over the winter and spring. All wood trim inthe house was replaced, new windows were installedwhere appropriate, the kitchen and upstairs bath-room were completely renovated, and the entire inte-rior was painted. Several laboratories received newhigh quality countertops along with other minorupgrades.

Additional paving of both driveways to ShalerHall was completed, including the parking lot for the Harvard Forest fleet, which made snow removalmuch easier. Lastly, a new Ford F-550 dump truckwas purchased to help in woods road improvementefforts and to provide a versatile vehicle for otheruses.

A new garage was started along Propsect HillRoad and behind Fisher House. The foundation andslab were excavated and poured and the 30' x 40'two-bay vehicle maintenance facility should be com-pleted by spring of 2004. This new building will con-tain proper facilities for storing hazardous waste inour ongoing effort to minimize Harvard Forest’senvironmental impact. Most importantly the newgarage will greatly improve our capacity to maintainour growing fleet of vehicles and equipment.

In addition significant improvements were madeto the kitchen in Shaler Hall and to the groundsaround most of the residential, office, and laboratorybuildings.

LIBRARY AND ARCHIVES

A Library Committee consisting of Edythe Ellin,John Burk, Glenn Motzkin, Emery Boose, andBarbara Flye met regularly to discuss and plan theshort- and long-term goals of the library. In June,work began on a reorganization project seeking tomaximize the space available in the current libraryand common room areas. Julie Pallant and KelliGraves have been working to update and consolidatethe numerous library databases.

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Site preparation for the new garage

Oscar Lacwasan

In the Archives, improvements included the addi-tion of a new map cabinet, a large new shelving unitfor the sample archives, and reorganization of the his-torical and photographic collections. John Burk andBarry Tomlinson sorted out a large collection ofassorted material from the past research of Drs.Tomlinson, John Torrey, and Martin Zimmermann.

INFORMATION MANAGEMENT ANDTECHNOLOGY ADVANCEMENT

The Harvard Forest purchased and installed a com-prehensive backup software system that automatical-ly captures PC data and safely stores it on our localservers. Since approximately ninety percent of ourcritical information resides on individual desktops,this new system provides us with a critical level ofdata protection that has not been available in the past.

Harvard Forest Web Site

The Harvard Forest Web site underwent a completeredesign this year and was launched this June. Thenew site features greatly expanded content, a friend-lier and more engaging interface, and beautiful pic-tures. The redesign includes a new name and can beseen at http://harvardforest.fas.harvard.edu. In addi-tion, the launch was concurrent with the move of theWeb server and site administration to the HarvardForest from its previous location at the LTER net-work office in Albuquerque.

Fisher Museum Web Site

The Fisher Museum at Harvard Forest received anew and improved Web page this year, http://harvardforest.fas.harvard.edu/museum.html. The site,designed by Julie Pallant, features for the first timethe dioramas online. A picture of each diorama isaccompanied by a text description. A greater audi-ence can be reached with the advent of the Web andif it is not possible to travel to the Harvard Forest toenjoy the dioramas, the on-line feature will providevisitors an appreciation for their workmanship andeducational value. In addition, a virtual tour of theSanderson Farm Natural History Interpretive Trailwas added to the Web site: http://harvardforest.fas.harvard.edu/museum/sanderson/sanderson.html.This outreach opportunity coincides with the instal-lation of the new interpretive signs along the trailitself.

Information Management

The Forest’s online Data Archive (http://harvard forest.fas.harvard.edu/data/archive.html) was re-designed and expanded by Emery Boose over thepast year in preparation for the LTER Site Review inJuly 2003. Metadata (documentation) for all 105 proj-ects currently posted on the Web was updated andconverted to Ecological Metadata Language (EML).A subset of XML (Extensible Markup Language)optimized for use in ecology, EML was developed by researchers in the LTER program and at theNational Center for Ecological Analysis andSynthesis (NCEAS). Adopted by the LTER Networkas its new standard for scientific metadata, EML willgreatly enhance the ability to locate, interpret, andintegrate scientific datasets.

ACTIVITIES OF THE HARVARD FORESTSTAFF

Audrey Barker Plotkin coordinated a field tour ofHarvard Forest for the Fifty-fifth Annual North-eastern Forest Soils Conference. The Conference hadheld its first meeting at Harvard Forest in 1939.Audrey also attended a Society of American Forestersworkshop, “Advances in Forest Health.”

Emery Boose and David Foster joined the newlyformed LTER Network Information System Ad-visory Group (NISAG), which brings together infor-mation managers and principal investigators and willadvise the LTER Network in the development of astrategic plan for information management and tech-

36

nology. Emery attended the LTER InformationMangers meeting in Orlando, the InformationMangers Executive Committee (IMEXEC) meetingin Santa Barbara, and the NISAG and LTERCoordinating Committee meetings at the KelloggBiological Station in Michigan.

John Burk gave natural history presentations atseveral institutions, including the Millers RiverEnvironmental Center, the Maine Audubon Society,and the Wildlife Learning Exchange in Concord. Hecontinued as a regular contributor to Natural NewEngland and Worcester County Online magazines, and asa participant in the state Biodiversity Days program.

Aaron Ellison presented seminars on his researchon pitcher plant communities at Harvard University,the University of Pennsylvania, Worcester StateCollege, and to the Concord Land Trust. New statis-tical models of pitcher plant demography were pre-sented at the Ecological Society of America (ESA)meetings in Savannah. He also participated as the“ant expert” in BioBlitzes at Tully Mountain andAcadia National Park. Aaron presented summaries ofhis research on ant community structure at aSymposium on Invertebrate Conservation in NewEngland sponsored by the American Museum of

Natural History, and at the annual meeting of theAcadian Entomological Society. He continued hisstints as associate editor-in-chief for Ecology andEcological Monographs and as associate editor forAmerican Journal of Botany. Aaron represented theForest at the LTER Coordinating Committee meet-ing at the Kellogg Biological Station, headed up anexternal review panel for the biology department atFranklin and Marshall College, and was a judge atthe western Massachusetts regional science fair. Heled field trips to Hawley Bog for the HitchcockCenter for the Environment, and to Mt. Tom for theNew England Wild Flower Society. Closer to homeAaron began a three-year term on the RoyalstonConservation Commission, and serves on the LandProtection Committee of the Mt. Grace Land Trust.

David Foster led the Harvard Forest FreshmanSeminar in the spring and Biology 299, Forest Practiceand Research, Ernie Gould’s original course in the falland spring. He participated in a symposium at theAmerican Society of Environmental History inProvidence that recognized and discussed the twenti-eth anniversary of Bill Cronon’s book on NewEngland landscape history, Changes in the Land. Davidjoined the advisory board for the ConservationInstitute of the Trustees of Reservations and he con-tinues to serve on boards and committees for theConservation Research Foundation, HighsteadArboretum, and Harvard University’s DavidRockefeller Center for Latin American Studies andCenter for Health and the Global Environment.David gave a keynote talk on New England’s ecolog-ical history at a USDA symposium at the Universityof Connecticut, an evening gallery talk on the eco-logical interpretation of Connecticut Valley land-scape painting at Mount Holyoke College, and invit-ed lectures at the University of New Hampshire andGeorge Perkins Marsh Institute at Clark University.He participated in a symposium of LandscapeLegacies at the Sevilletta LTER site in Albuquerque,the Ag Trans Biocomplexity workshop at ArizonaState University, and the LTER CoordinatingCommittee meeting at Niwot Ridge in Colorado.David and John Aber finished editing the HarvardForest LTER volume, Forests in Time — Forest Structureand Function as a Consequence of 1000 years of Change inNew England, and delivered the manuscript to YaleUniversity Press. In August David and his familyjoined the Harvard Alumni Association on its Baltictrip and in January he led a group from the HarvardNatural History Museums through Chilean and

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Aaron Ellison

Argentine Patagonia. Julian Hadley presented research results at the

Ameriflux meeting sponsored by the U.S. Depart-ment of Energy in October 2002 and at an AmericanGeophysical Union (AGU) meeting in December2002.

Brian Hall attended the New England Arc UsersConference, Bretton Woods, N.H., and HarvardUniversity GIS Day Symposium, presenting a poster“300 Years of Forest Change in Massachusetts: ForestCover in 1830.”

Holly Jensen-Herrin presented a poster at theNorth American Benthological Society annual meet-ing in Athens, Georgia, summarizing the preliminaryresults for her master’s thesis on stream biota in rela-tion to hydrology and other habitat variables in aseries of streams at Mt. Wachusett.

Dave Kittredge and David Foster attended theinitial meeting of the NSF biocomplexity project onTransitions in Agrarian landscapes in Phoenix inDecember. This LTER project seeks to identifymulti-scale patterns in agrarian landscape dynamics,and the ecological implications of this history forthose altered regions.

Dana MacDonald led two field trips for theUniversity of Massachusetts Department of Land-scape Architecture and Regional Planning, and oneclass field trip at the Harvard Forest for students ofthe Bazzaz Lab at Harvard University. He attendedthe Pine Barrens Research Forum at BrookhavenNational Laboratories, Brookhaven, N.Y., and co-leda one-day Harvard Forest field trip for students of theUniversity of Quebec, Montreal. Dana worked as a

volunteer to plan and site a long-term deer browsingexperiment at the Nature Conservancy’s Mash-omack Preserve on Shelter Island, N.Y., worked on acollaborative project with Dr. Alison Berry ofUniversity of California at Davis coring Uncas Pond,Mass. and subsequent sub-sampling of sediment, andco-organized the weekly seminar series at theHarvard Forest with Audrey Barker Plotkin.

Glenn Motzkin and Dave Orwig presented theirresearch on ridgetop communities at the annualEcological Society of America meeting in Tuscon.Glenn received the Massachusetts State Award fromthe New England Wildflower Society, and DavidFoster, Glenn, and Ben Slater were awarded the W. S.Cooper Award for 2003 by the Ecological Society ofAmerica. Glenn continues to serve as an EcologyAdvisor for the Trustees of Reservations, and anassociate member of the Massachusetts NaturalHeritage and Endangered Species Program AdvisoryCommittee. He served on the thesis committee ofSally Shaw from UMASS, and served as a critic forthe Conway School of Landscape Design (CSLD)graduate projects. Glenn led several field trips toMontague Plain, including one with Tim Simmons ofthe Massachusetts Natural Heritage and EndangeredSpecies Program for the Town of Montague as part ofthe annual Biodiversity Days.

Sylvia Barry Musielewicz presented a poster atthe International Congress of Limnogeology inTucson, “2000 years of climate change from pollenrecords in Massachusetts.”

John O’Keefe gave talks on Harvard Forestresearch and the history of northeastern forests at theUniversity of Massachusetts, the Squam LakeScience Center in Holderness, N.H., and to the

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John Wisnewski

Harvard Forest Friends Meeting — 2002

Annual Meeting of the Rhode Island Wild PlantSociety. He attended the Agricultural Transitions/Biocomplexity workshop in Phoenix in December asHarvard Forest’s education representative, theAmerican Society for Environmental History/ForestHistory Society meeting in Providence, R.I., inMarch, and the first-ever International ForestryMuseums meeting at the World Forestry Center inPortland, Oreg., in May. John serves on the boards ofthe Mount Grace Land Conservation Trust, wherehe is vice-president, Massachusetts Forestry Asso-ciation and Millers River Environmental Center, andon the executive committee of the North QuabbinRegional Landscape Partnership. He also continuesto serve on the Quabbin Science and TechnicalAdvisory Committee and the EnvironmentalSecretary’s Advisory Group on EnvironmentalEducation.

Dave Orwig presented talks at HampshireCollege, the Notch Visitor Center for the Massa-chusetts Department of Environmental Management,the Advances in Forest Health Workshop, and theNorth American Forest Ecology Workshop inCorvallis, Oreg. He presented a poster at theEcological Society of America Annual Meeting inTucson, Ariz. Dave served on the thesis committees

of Aaron Kimple at Bard College, Sarah Parker atAntioch College, and is the Ph.D. dissertation advi-sor to former Harvard Forest REU student, AnthonyD’Amato at the University of Massachusetts. He ledfield trips at the Quabbin Reservoir and HarvardForest for the Freshman Seminar Class.

Barry Tomlinson attended the Annual Meetingof the Botanical Society of America at the Universityof Wisconsin in Madison in early August presentingresearch on Gnetum. In April he attended the third ofthe recurrent meetings on the systematics, evolution,and biology of monocotyledons, this time sponsoredby Rancho Santa Ana Botanic Garden, Claremont,California, held at the Ontario Convention Center.The winter months were spent at the NationalTropical Botanical Gardens headquarters on Kauai,Hawaii, with several visits to their outstation at “TheKampong,” Coconut Grove, Miami, Florida, withcontinued collaborative work with staff of theFairchild Tropical Garden, Coral Gables, Florida.

In June, Barry served on a panel at theUniversity of Miami, reviewing the place in the uni-versity’s teaching, research, and outreach programsof the rich campus collections of tropical plants thatmake this university unique in the continental UnitedStates.

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Welcome barbecue for the summer students

John Aber University of New HampshireMark Ashton Yale UniversityFakhri Bazzaz Harvard UniversityRichard Bowden Allegheny College Frank Bowles Ecosystems Center — MBLAlfram Bright Harvard UniversityRobert Brooks USDA Forest ServiceDavid Bryant Harvard UniversityJohn Budney Harvard UniversityLiz Burroughs Ecosystems Center — MBLElizabeth Chilton University of MassachusettsBrendan Choat Harvard UniversityV. Y. Chow Harvard UniversityScott Costa University of VermontPatrick Crill University of New HampshireWilliam Currie University of MichiganEric Davidson Woods Hole Research CenterKathleen Donohue Harvard UniversityJim Ehleringer University of UtahDavid Fizjarrald SUNY, AlbanyJulia Gaudinski UCLA, IrvineElaine Gottleib Harvard UniversityN. Michele Holbrook Harvard UniversityDavid Hollinger USDA Forest ServiceLucy Hutyra Harvard UniversityChristine Jones Harvard UniversitySylvan Kaufman Harvard Unversity

Alan Kirschbaum University of WisconsinChun-Ta Lai University of UtahCathy Langtimm Holy Cross CollegeKristin Lewis Harvard UniversityAlison Magill Univ. of New HampshireLynn Margulis University of MassachusettsMary Martin Univ. of New HampshireJerry Melillo Ecosystems Center — MBLPatricia Micks Ecosystems Center — MBLJacqueline Mohan Duke UniversityJeff Morisette NASASarah Morisseau Ecosystems Center — MBLMitch Mulholland University of MassachusettsRanga Myneni Boston UniversityJ. William Munger Harvard UniversityKnute Nadelhoffer University of MichiganTommi Nyman Harvard UniversityJennifer Pontius U.S. Forest ServiceElizabeth Pyle Harvard UniversityKathleen Savage Woods Hole Research CenterTim Sipe Franklin & Marshall CollegeRolf Staebler SUNY, AlbanyPaul Steudler Ecosystems Center — MBLSusan Trumbore University of CaliforniaShawn Urbanski Harvard UniversityMaciej Zwienecki Harvard University

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VISITING RESEARCH SCIENTISTS AT THE HARVARD FOREST 2002–2003

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

Fisher House

PUBLICATIONS

Anderson, R. L. 2001. Integrating lateral expansioninto models of peatland development in temper-ate New England. M.F.S. thesis, HarvardUniversity.

Anderson, R. L., D. R. Foster, and G. Motzkin. 2003.Integrating lateral expansion into models of peat-land development in temperate New England.Journal of Ecology 91: 68–76.

Bellemare, J., G. Motzkin, and D. R. Foster. 2002.Legacies of the agricultural past in the forestedpresent: an assessment of historical land-useeffects on rich mesic forests. Journal ofBiogeography 29: 1401–1420.

Berlik, M. M., D. B. Kittredge, and D. R. Foster.2002. The illusion of preservation: a global envi-ronmental argument for the local production ofnatural resources. Harvard Forest Paper No. 26.

Berlik, M. M., D. B. Kittredge and D. R. Foster.2002. The illusion of preservation: a global envi-ronmental argument for the local production ofnatural resources. Journal of Biogeography 29:1557–1568.

Catovsky, S. and F. A. Bazzaz. 2002. Feedbacksbetween canopy composition and seedling regen-eration in mixed conifer broad-leaved forests.Oikos 98: 403–420.

Catovsky, S. and F. A. Bazzaz. 2002. Nitrogen avail-ability influences regeneration of coniferous andbroad-leaved tree species in the understoryseedling bank. Ecological Applications 12:1056–1070.

Catovsky, S. and F. A. Bazzaz. 2002. Plant competi-tion in an elevated CO2 world. Pp. 471–481 InH. A. Mooney and J. Canadell (Eds.), Encyclopediaof Global Change. John Wiley and Sons, Ltd.,Chichester, England.

Catovsky, S., N. M. Holbrook, and F. A. Bazzaz.2002. Coupling whole-tree transpiration andcanopy photosynthesis in coniferous and broad-leaved tree species. Canadian Journal of ForestScience 32: 295–309.

Cobb, R. C. and D. A. Orwig. 2002. Impact of hem-lock woolly adelgid on decomposition: anoverview. Pp. 317–322 In R. C. Reardon, B. P.Onken, and J. Lashomb (Eds.), Symposium on theHemlock Woolly Adelgid in the Eastern United StatesProceedings. New Jersey Agricultural ExperimentStation Publication, New Brunswick, N.J.

Cogbill, C. V., J. Burk, and G. Motzkin. 2002. The

forests of presettlement New England, U.S.A.:spatial and compositional patterns based on townproprietor surveys. Journal of Biogeography 29:1279–1304.

Davidson, E. A., K. Savage, L. V. Verchot, and R. I.Navarro. 2003. Minimizing artifacts and biases inchamber-based measurements of soil respiration.Agricultural and Forest Meteorology 113: 21–37.

Eberhardt, R., D. R. Foster, G. Motzkin, and B. Hall.2003. Conservation of changing landscapes: veg-etation, land-use history, and fire on Cape CodNational Seashore. Ecological Applications 13:68–84.

Ellison, A. M., E. J. Farnsworth, and N. J. Gotelli.2002. Ant diversity in pitcher plant bogs ofMassachusetts. Northeastern Naturalist 9: 267–284.

Ellison, A. M., N. J. Gotelli, J. S. Brewer, L. Cochran-Stafira, J. Kneitel, T. E. Miller, A. S. Worley, andR. Zamora. 2003. The evolutionary ecology ofcarnivorous plants. Advances in Ecological Research33: 1–74.

Field, T. S., P. J. Franks, and T. L. Sage 2003.Ecophysiological shade adaptation in the basalangiosperm, Austrobaileya scandens (Austro-baileyaceae). International Journal of Plant Sciences164: 313–324.

Finley, A. O. 2002. Forest owner attitudes towardsmanagement at a multi-property scale. M.S. the-sis, University of Massachusetts.

Forman, R. T. T., B. Reineking and A. M.Hersperger. 2002. Road traffic and nearby grass-land bird patterns in a suburbanizing landscape.Environmental Management 29: 782–800.

Forman, R. T. T., D. Sperling, J. A. Bissonette, A. P.

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Community House

Clevenger, C. D. Cutshall, V. H. Dale, L. Fahrig,R. France, C. R. Goldman, K. Heanue, J. A.Jones, F. J. Swanson, T. Turrentine, and T. C.Winter. 2003. Road Ecology: Science and Solutions.Island Press, Washington, D.C. 418 pp.

Foster, D. R. 2002. Insights from historical geogra-phy to ecology and conservation: lessons fromthe New England landscape. Journal ofBiogeography 29: 1269–1275.

Foster, D. R. 2002. Conservation issues andapproaches for dynamic cultural landscapes.Journal of Biogeography 29: 1533–1535.

Foster, D. R. 2002. Thoreau’s country: a historical-ecological approach to conservation of the NewEngland landscape. Journal of Biogeography 29:1537–1555.

Foster, D. R., G. Motzkin, D. Bernardos, and J.Cardoza. 2002. Wildlife dynamics in the chang-ing New England landscape. Journal ofBiogeography 29: 1337–1357.

Foster, D. R., S. Clayden, D. A. Orwig, B. Hall, andS. Barry. 2002. Oak, chestnut and fire: climaticand cultural controls of long-term forest dynam-ics in New England. Journal of Biogeography 29:1359–1379.

Foster, D. R., B. Hall, S. Barry, S. Clayden, and T.Parshall. 2002. Cultural, environmental, and his-torical controls of vegetation patterns and themodern conservation setting on the island ofMartha’s Vineyard, U.S.A. Journal of Biogeography29: 1381–1400.

Foster, D. R., F. Swanson, J. Aber, I. Burke, N.Brokaw, D. Tilman and A. Knapp. 2003. Theimportance of land-use legacies to ecology andconservation. BioScience 53: 77–88.

Franks, P. J. 2003. Use of the pressure probe in stud-ies of stomatal function. Journal of ExperimentalBotany 54: 1–10.

Gerhardt, F. and D. R. Foster. 2002. Physiographicaland historical effects on forest vegetation in cen-tral New England, U.S.A. Journal of Biogeography29: 1421–1437.

Gotelli, N. J. and A. M. Ellison. 2002. Assembly rulesfor New England ant assemblages. Oikos 99:591–599.

Gotelli, N. J. and A. M. Ellison. 2002. Nitrogen dep-osition and extinction risk in the northern pitch-er plant Sarracenia purpurea. Ecology 83: 2758–2765.

Hall, B., G. Motzkin, D. R. Foster, M. Syfert, and J.Burk. 2002. Three hundred years of forest andland-use change in Massachusetts, U.S.A. Journal

of Biogeography 29: 1319–1335. Hersperger, A. M. and R. T. T. Forman. 2003.

Adjacency arrangement effects on plant diversityand composition in woodland patches. Oikos 101:279–290.

Holbrook, N. M., M. A. Zwieniecki and P. J.Melcher. 2002. The dynamics of “dead” wood:maintenance of water transport through plantstems. Integrative and Comparative Biology 42:492–496.

Kizlinski, M., D. A. Orwig, R. C. Cobb, and D. R.Foster. 2002. Direct and indirect ecosystem con-sequences of an invasive pest on forests dominat-ed by eastern hemlock. Journal of Biogeography 29:1489–1503.

Lawrence, D. and D. R. Foster. 2002. Changes in

forest biomass, litter dynamics and soils follow-ing shifting cultivation in southern Mexico: anoverview. Interciencia 27: 400–408.

Melillo, J. M., P. A. Steudler, J. D. Aber, K. Newkirk,H. Lux, F. P. Bowles, C. Catricala, A. Magill, T.Ahrens. and S. Morrisseau. 2002. Soil warmingand carbon-cycle feedbacks to the climate sys-tem. Science 298: 2173–2176.

Motzkin, G. and D. R. Foster. 2002. Grasslands,heathlands, and shrublands in New England: his-torical interpretation and approaches to conser-vation. Journal of Biogeography 29: 1569–1590.

Motzkin, G., S. C. Ciccarello, and D. R. Foster. 2002.Frost pockets on a level sand plain: does varia-tion in microclimate help maintain persistent veg-

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Higginson House

etation patterns? Journal of the Torrey BotanicalSociety 129: 154–163.

Motzkin, G., D. A. Orwig, and D. R. Foster. 2002.Vegetation and disturbance history of a raredwarf pitch pine community in western NewEngland, U.S.A. Journal of Biogeography 29:1455–1467.

Motzkin, G. M., R. Eberhardt, B. Hall, D. R. Foster,J. Harrod, and D. MacDonald. 2002. Vegetationvariation across Cape Cod, Massachusetts: envi-ronmental and historical determinants. Journal ofBiogeography 29: 1439–1454.

Muth, C. C. and F. A. Bazzaz. 2002. Tree canopy dis-placement at forest gap edges. Canadian Journal ofResearch 32: 247–254.

Muth, C. C. and F. A. Bazzaz. 2002. Tree seedlingcanopy responses to conflicting photosensorycues. Oecologia 132: 197–204.

Ollinger, S., J. D. Aber, P. B. Reich, and R. J. Freuder.2002. Interactive effects of nitrogen deposition,tropospheric ozone, elevated CO2 and land usehistory on the carbon dynamics of northernhardwood forests. Global Change Biology 8:545–562.

Orwig, D. A. 2002. Ecosystem to regional impacts ofintroduced pests and pathogens: historical con-text, questions and issues. Journal of Biogeography29: 1471–1474.

Orwig, D. A. 2002. Stand dynamics associated withchronic hemlock woolly adelgid infestations insouthern New England. Pp. 36–47 In R. C.Reardon, B. P. Onken, and J. Lashomb (Eds.),Symposium on the Hemlock Woolly Adelgid in EasternNorth America Proceedings. New Jersey AgriculturalExperiment Station Publication, New Brunswick,N.J.

Orwig, D. A., D. R. Foster, and D. L. Mausel. 2002.Landscape patterns of hemlock decline in NewEngland due to the introduced hemlock woollyadelgid. Journal of Biogeography 29: 1475–1487.

Orwig, D. A. and M. L. Kizlinski. 2002. Vegetationresponse following hemlock woolly adelgid infes-tation, hemlock decline, and hemlock salvagelogging. Pp. 106–117 In R. C. Reardon, B. P.Onken, and J. Lashomb (Eds.), Symposium on theHemlock Woolly Adelgid in the Eastern United StatesProceedings. New Jersey Agricultural ExperimentStation Publication, New Brunswick, NJ.

Paillet, F. L. 2002. Chestnut: history and ecology ofa transformed species. Journal of Biogeography 29:1517–1530.

Parshall, T. and D. R. Foster. 2002. Fire on the NewEngland landscape: regional and temporal varia-tion, cultural and environmental controls. Journalof Biogeography 29: 1305–1317.

Parshall, T., D. R. Foster, E. Faison, D. MacDonald,and B. C. S. Hansen. 2003. Long–term history ofvegetation and fire in pitch pine-oak forests onCape Cod, Massachusetts. Ecology 84: 736–738.

Posluszny, U. and P. B. Tomlinson. 2003. Aspects ofinflorescence and floral development in the puta-tive basal angiosperm Amborella trichopoda(Amborellaceae). Canadian Journal of Botany 81:28–39.

Sack, L., P. D. Cowan, and N. M. Holbrook. 2003.The major veins of mesomorphic leaves, revisit-ed: a test of conductive overload in leaves of Acersaccharum (Aceraceae) and Quercus rubra(Fagaceae). American Journal of Botany 90: 32–39.

Savage, K. E. and E. A. Davidson. 2003. A compari-son of manual and automated systems for soilCO2 flux measurements: trade-offs between spa-tial and temporal resolution. Journal ofExperimental Botany 54: 891–899.

Tingley, M. W., D. A. Orwig, R. Field, and G.

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Motzkin. 2002. Avian response to removal of aforest dominant: consequences of hemlock wool-ly adelgid infestations. Journal of Biogeography 29:1505–1516.

Tomlinson, P. B. 2003. Development of gelatinous(reaction) fibers in stems of Gnetum gnemon(Gnetales). American Journal of Botany 90: 965–972.

Tomlinson, P. B. and T. Takaso. 2002. Seed conestructure in conifers in relation to developmentand pollination: a biological approach. CanadianJournal of Botany 80: 1250–1273.

Tomlinson, P. B. and M. H. Zimmermann. 2003.Stem vascular architecture in the Americanclimbing palm Desmoncus (Arecaceae-Arecoideae-Bactridinae). Botanical Journal of the Linnean Society142: 243–254.

Turner, B. L. II, S. C. Villar, D. R. Foster, J.Geoghegan, E. Keys, P. Klepeis, D. Lawrence, P.M. Mendoza, S. Manson, Y. Ogneva-Himmelberger, A. B. Plotkin, D. P. Salicrup, R.R. Chowdhury, B. Savitsky, L. Schneider, B.Schmook, and C. Vance. 2001. Deforestation inthe southern Yucatan peninsular region: an inte-grative approach. Forest Ecology and Management154: 353–370.

Tyree, M. T. and M. H. Zimmermann. 2003. Xylemstructure and the ascent of sap. Springer Verlag,Berlin. [2nd edition]

Woodcock, D. W. and A. D. Shier. 2002. Wood spe-cific gravity and its radial variations: the manyways to make a tree. Trees 16: 437–443.

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Walk-up tower for sampling gas exchange in the Prospect Hill hemlock stand

ACKNOWLEDGMENT OF SUPPORT

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

Friends of the Harvard Forest Andrew W. Mellon FoundationCommonwealth of Massachusetts

Department of Environmental ManagementDivision of Fisheries and Wildlife Natural Heritage & Endangered Species

ProgramDepartment of Energy, National Institute for

Global Environmental ChangeHarvard University

Center on the EnvironmentDavid Rockefeller Center

Massachusetts Environmental TrustNational Tropical Botanical GardenNational Science Foundation

Digital Government ProgramEcosystem StudiesInternational ProgramsInformation Technology Research Long-term StudiesEcology ProgramResearch Experience for Undergraduates

Southern Taconics Research and Conservation Center

Sweetwater TrustThe Nature Conservancy

GIFTS

The Harvard Forest Library and Archives thanks thefollowing for gifts received this past year: RichardForman for his book Road Ecology: Science and Solutions,Jack Putz for Biodiversity Conservation in the Context ofTropical Forest Management, John Woolsey for NaturalAreas Journal, Bob Howarth for The Nitrogen Cycle atRegional to Global Scales, James Baird for numerousjournals, Mel Tyree for the second edition of XylemStructure and the Ascent of Sap, an update of the first edi-tion written by Martin Zimmermann in 1984, andMichael Caduto for A Time Before New Hampshire: TheStory of a Land and Native Peoples, which included oneof Hugh Raup’s historical photos. David Tatlockadded numerous historical documents to the R. T.Fisher collection, and Michael Scott loaned a copy ofthe 1898 Atlas of Worcester County.

NEW FUNDING

Aaron Ellison, Nick Gotelli, Nate Sanders, and ErikJules. Community re-assembly after a catastrophicfire. Collaborative with Humboldt State Universityand the University of Vermont. National ScienceFoundation. $8,000 sub-award (of $50,000 award).

Aaron Ellison and Nick Gotelli. Effects of nutrientstress on a co-evolved food web. Collaborative withUniversity of Vermont. National Science Foundation.$323,466 primary award (of $350,000 total award).

David Foster and Elizabeth Colburn. Establishing along-term monitoring protocol for freshwater macro-invertebrates at Cape Cod National Seashore.National Park Service. $51,001.

David Foster, Elizabeth Chilton, Glenn Motzkin, andothers. Agrarian landscapes in transition: a cross-scale approach. National Science Foundation. $228,672. (Subaward with Arizona State University).

David Foster, Ana Barros, Kathleen Donohue, AaronEllison, Henry Foster, Michelle Holbrook, JamesLevitt, and Paul Moorcroft. Ecological and environ-mental impacts of the extinction of core species.Harvard University Center for the Environment.$46,000.

David Foster, Betsy Colburn, Emery Boose, and oth-ers. Development of a hydrological infrastructure forheadwater streams at the Harvard Forest. NationalScience Foundation, Long Term Ecological Researchprogram. $ 63,000

David Foster. Ecological history and conservation ofthe southern Patagonian landscape. David Rocke-feller Center for Latin American Studies at HarvardUniversity. $8,000.

Julian Hadley and David Foster. Evaluating theeffects of diverse vegetation types and soil drainageclasses on net carbon exchange of a landscape mosa-ic with mobile and fixed eddy covariance systems.Department of Energy National Institute for GlobalEnvironmental Change. $125,000.

Glenn Motzkin, David Foster, and Dave Kittredge.Ecological and conservation implications of forestharvesting patterns across Massachusetts. R. J. Kose

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Award. The Nature Conservancy. $100,000.

Dave Orwig, David Kittredge, and David Foster.Forest response to the decline of a dominant species:ecosystem to regional analyses of the impact of thehemlock woolly adelgid on northeastern forests.National Science Foundation. $400,000. In addition,a $12,000 supplemental grant from NSF to fund anREU summer student for this project.

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David R. Foster Petersham, MassachusettsDirector August 2003