THE HARVARD FOREST 2004–2006

Harvard University

Front Cover: Misty Morning on Prospect Hill, John BurkBack Cover: Pasture at Harvard Forest, Tracy RogersPhotography: Audrey Barker Plotkin, Emery Boose, Laura Briscoe, Brian DeGasperis, John Burk, Hilary

Crowell, Stephanie Day, John Klironomos, Tracy Rogers, Brynne Simmons, Pamela Snow,Jimmy Tran, and Ben Wolfe.

Clearing of a homestead by an early settler A.D. 1740. Harvard Forest Diorama.

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ANNUAL REPORT OF THE HARVARD FOREST 2004–2006

Contents

Personnel at the Harvard Forest - - - - - - - 2Introduction to the Harvard Forest - - - - - - - 3Transitions and Accolades - - - - - - - - 3New Staff - - - - - - - - - - 4Research Activities - - - - - - - - - 4Bullard Fellows - - - - - - - - - 30Educational Activities - - - - - - - 33

Summer Research Program - - - - - - - 33Master of Forest Science Program - - - - - - 36Building the LTER Schoolyard Program - - - - - 36

Activities of the Fisher Museum - - - - - - - 37Meetings, Conferences, Seminars, 2004–2005 - - - - - 38Meetings, Conferences, Seminars, 2005–2006 - - - - - 38

Forest Management and Infrastructure - - - - - - 41Library and Archives - - - - - - - - 41Information Management and Technology Advancement - - - - 42Activities of the Harvard Forest Staff - - - - - - 42Land Protection Efforts - - - - - - - - 45Visiting Research Scientists at the Harvard Forest 2004–2005 - - - 47Visiting Reserach Scientists at the Harvard Forest 2005–2006 - - - 49Publications - - - - - - - - - 51Acknowledgment of Support and Gifts - - - - - - 58New Funding 2005 - - - - - - - - - 59New Funding 2006 - - - - - - - - - 60

http://harvardforest.fas.harvard.edu

Audrey Barker Plotkin Research AssistantEmery Boose Information & Computer

System ManagerJeannette Bowlen AccountantJohn Burk Archivist & Librarian Posy Busby MFS StudentJessica Butler Research AssistantLaurie Chiasson Receptionist/Accounting

AssistantElizabeth Colburn Aquatic EcologistEowyn Connolly-Brown Laboratory TechnicianAnthony D’Amato Graduate StudentBrian DeGasparis MFS StudentBrian Donahue Environmental HistorianElaine Doughty Research AssistantEdythe Ellin Director of AdministrationEd Faison MFS StudentChristian Foster Laboratory TechnicianDavid Foster DirectorLucas Griffith Maintenance TechnicianJulian Hadley EcophysiologistBrian Hall Research AssistantLinda Hampson Staff AssistantAmber Jarvenpaa Assistant Summer CookJim Karagatzides Post-doctoral FellowDavid Kittredge Forest Policy AnalystPaul Kuzeja Research AssistantOscar Lacwasan Building ServicesJames Levitt Director, Program on

Conservation InnovationHeidi Lux Research AssistantBrooks Mathewson MFS StudentRobert McDonald Post-doctoral FellowJason McLachlan Post-doctoral FellowJacqueline Mohan Post-doctoral FellowGlenn Motzkin Plant EcologistJohn O’Keefe Museum & Schoolyard

CoordinatorDavid Orwig Forest EcologistWyatt Oswald Paleoecologist Julie Pallant System & Web

AdministratorRichard Schulhof MFS StudentMichael Scott Maintenance TechnicianJudy Shaw Building ServicesPamela Snow Environmental EducatorBill Sobczak Aquatic EcologistKristina Stinson Research AssociateTravis Stolgitis Maintenance TechnicianP. Barry Tomlinson E. C. Jeffrey Professor of

Biology, EmeritusBetsy Von Holle Post-doctoral FellowJohn Wisnewski Maintenance TechnicianTim Zima Summer Cook

Bullard Fellows

John Briggs Arizona State UniversityRichard Bowden Allegheny CollegeElizabeth Farnsworth N. E. Wild Flower SocietyMark Harmon Oregon State UniversityJulia Jones Oregon State UniversityGreg Jordan University of Tasmania–

HobartDavid Lee Florida International U.Gidon Ne’eman University of Haifa-Oranim,

IsraelThomas Sinclair University of Florida Conghe Song University of North

Carolina at Chapel HillPeter Thomas Keele University, UKJill Thompson University of Puerto RicoJohn Weishampel University of Central Florida

Harvard University Affiliates

Alfram Bright DEAS*John Budney DEAS*Brendan Choat OEB*Daniel Curran DEAS*Bruce Daube DEAS*Peter del Tredici Arnold ArboretumDavid Diaz Harvard CollegeKathleen Donohue OEB*Richard Forman Graduate School of DesignCharles H.W. Foster JFK School of Government Elaine Gottleib DEAS*N. Michelle Holbrook OEB*Lucy Hutyra EPS*Kathryn McKain DEAS*David Medvigy DEAS*Paul Moorcroft OEB*Michael Muilenberg School of Public HealthJ. William Munger EPS*Elizabeth Pyle DEAS*Fulton Rockwell OEB*Christine Rogers School of Public HealthSanna Sevanto OEB*Rachel Spicer OEB*Steve Wofsy DEAS*Maciej Zwieniecki Arnold Arboretum

*Museum of Comparative Zoology Organismic & Evolutionary BiologyEarth & Planetary SciencesDivision of Engineering & Applied Sciences

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PERSONNEL AT THE HARVARD FOREST 2004-2006

INTRODUCTION TO THE HARVARD FOREST

Since its establishment in 1907, the Harvard Foresthas served as Harvard University’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 town of Petersham, Massachusetts, thatinclude mixed hardwood and conifer forests, ponds,streams, extensive spruce and maple swamps, fields,and plantations. Additional land holdings include the25-acre Pisgah Forest in southwestern New Hamp-shire (located in the 13,500-acre Pisgah State Park), avirgin forest of white pine and hemlock that was 300years old when it blew down in the 1938 hurricane;the 100-acre Matthews Plantation in Hamilton, Mass-achusetts, which is largely composed of plantationsand upland forest; and the 90-acre Tall Timbers For-est in Royalston, Massachusetts. In Petersham, acomplex of buildings that includes Shaler Hall, theFisher Museum, and the John G. Torrey Laborato-ries provide office and laboratory space, computerand greenhouse facilities, and lecture rooms for sem-inars and conferences. Nine additional houses pro-vide accommodations for staff, visiting researchers,and students. Extensive records, including long-termdata sets, historical information, original field notes,maps, photographic collections, and electronic dataare maintained in the Harvard Forest Archives.

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

Forest 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 and Plane-tary Sciences (EPS), the School of Public Health(SPH), and the Graduate School of Design (GSD) atHarvard, and with the Department of NaturalResources Conservation at the University of Massa-chusetts, the Ecosystems Center of the Marine Bio-logical Laboratory, and the Complex SystemsResearch Center at the University of New Hampshire.

The staff and visiting faculty of approximatelyfifty work collaboratively to achieve the research, edu-cational, and management objectives of the Harvard Forest. A management group meets monthlyto discuss current activities and to plan future programs. Regular meetings with the HF LTER science team, weekly research seminars and lab dis-cussions, and an annual ecology symposium providefor an infusion of outside perspectives. The seven-member facilities crew undertakes forest managementand physical plant activities. The coordinator of theFisher Museum oversees many educational and out-reach programs.

Funding for the Harvard Forest is derived fromendowments, whereas major research support comesprimarily from the National Science Foundation,Department of Energy, U.S. Department of Agricul-ture, Andrew W. Mellon Foundation, and othergranting sources. Our Summer Program for StudentResearch is supported by the National Science Foun-dation, the A. W. Mellon Foundation and the R. T.Fisher Fund.

TRANSITIONS AND ACCOLADES

P. Barry Tomlinson was one of a select group ofprominent botanists who were awarded the Centen-nial Medal of the Botanical Society of America tocommemorate the one hundredth anniversary of thefounding of the Society in 1906.

Two Forest post-doctoral fellows moved ontonew positions. Robert McDonald is now beginning aone-year David H. Smith Conservation Research Fel-lowship at the Harvard University Graduate Schoolof Design. The fellowship which is a program of theSociety for Conservation Biology is devoted exclu-sively to applied conservation research problems.Rob will work with Richard Forman of the GSD andDr. Peter Kareiva, from The Nature Conservancy.

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Michael Bank, a post-doctoral fellow at the Forest fora year has moved on to be Research Fellow at theDepartment of Environmental Health at the HarvardSchool of Public Health. His research focuses onamphibian declines and species loss. Mike worked aspart of the research team investigating the effects offorest logging on vegetation patterns. Severalresearch assistants also departed in the past yearincluding: Sultana Jefts and Laura Pustell Barbash,both finished working at the Forest after two years asresearch assistants with the Hemlock Woolly Adelgidproject headed by David Orwig; Julie Hall whoworked on historical and spatial data analysis withGlenn Motzkin and David Foster; and Holly Jensen-Herrin who worked with Betsy Colburn on variousaquatic projects in Massachusetts.

NEW STAFF

Travis Stolgitis was hired as a new member of theWoods Crew. Travis’s specialty is carpentry and hehas extensive experience in post-and-beam construc-tion. Also, Eowyn Connolly-Brown, a junior at AtholHigh School, began working with the HWA researchteam led by David Orwig, assisting with lab and fieldwork.

RESEARCH ACTIVITIES

Flora of Harvard Forest

For nearly a century, Harvard Forest has been thefocus of a wide range of ecological investigations,with a strong emphasis on studies of vegetation, dis-turbance history, and stand dynamics. However,

there have been no previous attempts to evaluatechanges in the flora over time. In the early 1930s,Hugh Raup apparently developed the first checklistof vascular plants of Petersham, MA, based on hisextensive collections. Subsequently, C. Earle SmithJr. summarized the town’s vascular flora in 1947,based on the collections of Dr. I. M. Johnson as wellas other towns and a review of specimens in the Har-vard Forest and New England Botanical Clubherbaria. Over the past two years, Glenn Motzkin has workedclosely with Jerry Jenkins and Kirsten Ward to: 1)document the current flora including detailed infor-mation on the distribution and abundance of locallyand regionally uncommon species and invasivespecies; 2) evaluate changes in the flora over the pastcentury; 3) provide a strong baseline for document-ing future changes in the flora. This year, Glenn andJerry, working closely with Sue Williams, a bryolo-gist from Rowe, Massachusetts, who has extensiveexperience with the bryophytes (mosses and liver-worts) of the Northeast and summer researcherLaura Briscoe, are extending the flora project toinclude the bryophytes of Harvard Forest.

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Professor P. Barry Tomlinson.

Pitcher-Plant Ecology

Research on pitcher-plant ecology continued its pro-ductive trajectory as NSF renewed support for PIAaron Ellison’s work on this model ecological systemfor another five years. Aaron and research assistantJessica Butler focused the final year of their last granton completing a two-year experiment on nitrogenuptake, storage, and translocation by the northernpitcher plant, Sarracenia purpurea. The analysis of

pitchers and/or roots fed either 15N-enriched ammo-nium nitrate (to simulate acid rain) or pitchers fed15N-enriched fruit flies should allow us to finally con-struct a detailed nitrogen budget for this plant andcement its utility as a biological indicator for fine-scale assays of nitrogen deposition. As this report wasbeing written, the tissue analyses of the 15N-fed pitch-ers had just been completed by David Post’s lab atYale. Preliminary examination of the data suggeststhat microbial activity within the food web may dom-inate nitrogen cycling in this aquatic food web, incontrast to expectations based on many years ofresearch focusing on the role of the fly larvae at thetop of the pitcher’s aquatic food web.

The careful observations made in the summer of2005 by Bullard Fellow Gidon Ne’eman and his wifeRina Ne’eman (on sabbatical from Anne Frank HighSchool in Haifa) on pitcher-plant pollinationappeared November 2006 in the American Journal ofBotany. This summer, Jess Butler is monitoring plantsthat Gidi and Rina manipulated last year to deter-mine how variation in nutrient availability and pho-tosynthetic capacity in one year translate to changesin reproductive success in the subsequent year.NSERC post-doc Jim Karagatzides will be capitaliz-ing on these data to determine “construction costs”

(in terms of carbon) of various tissues of Sarracenia aswe probe more deeply into the evolutionary costs ofbotanical carnivory. Rina’s carefully collected set ofpollen of all ten species of Sarracenia growing in thegreenhouse were imaged and measured by ElaineDoughty and Wyatt Oswald in the Paleo Lab, and itappears that pollen of at least several species of Sar-racenia can be distinguished based on only two meas-urements. This work may lead to additional palyno-logical markers to identify patterns of herbaceousvegetation in many of the cores studied by the PaleoLab.

The focus for the next five years is on spatial rela-tionships among pitcher plants, ants (their primaryprey), and two species of little-studied noctuid moths,Exyra fax and Papaipema appassionata (the latter is aspecies of special concern in Massachusetts). Thesethree taxa — the ants, the plants, and the moths —make up the “terrestrial” food web of the Sarraceniasystem, and Aaron and Jess will be exploring howchanges in plant quality resulting from patterns ofmoth herbivory translate into changes in populationstructure of the ants, which not only are prey for theplants but also forage at the plants for sweet nectar.Some of the preliminary data that led to this success-ful proposal were collected by 2003 Research Experi-ence for Undergraduates (REU) student Dan Atwa-ter; his data were published in Northeastern Naturalist inspring 2006.

Aaron is also working to place this work in abroader historical context. In the early twentieth cen-tury, the Delaware entomologist and naturalist FrankMorton Jones did extensive work on Sarracenia andits insect associates throughout the eastern UnitedStates, as well as on the California pitcher plant, Dar-lingtonia californica. Jones’s archival papers from thisresearch, along with his entomological and botanicalcollections were donated to the Peabody Museum atYale. Aaron has the archival materials on long-termloan, and they are being digitized and compiled intoa Web site by 2006 REU student Stephanie Day.

Several exciting aspects stand out from this mate-rial. First, Jones carried on extensive correspondencewith major figures in nineteenth- and twentieth-century botany, including William Jepson and JohnMacFarlane, and compiled some of the key recordsof the collecting activities of Rebecca Austin (see illus-tration on next page), a California naturalist who madesome of the earliest and most detailed observationson the biology of Darlingtonia. Second, Jones’s workon Sarracenia and its prey in the southeastern U.S.

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predates the introduction of the red fire ant, Solenopsisinvicta, providing a window into how changes inspecies composition of insects in bogs may have ledto significant diet shifts of Sarracenia. Third, hisdetailed maps of localities will allow mapping of his-toric locations for Sarracenia and permit the assess-ment of the magnitude of region-wide declines in sev-eral species that are now federally endangered. Jonesalso did some of the only detailed work on bacterialpopulations within Sarracenia pitchers (published in1927!). Post-doc Celeste Petersen in OEB is workingon bringing bacterial studies of Sarracenia into themolecular age.

Jim Karagatzides’s post-doctoral research withDr. Aaron Ellison, funded by NSERC Canada, uses

the carnivorous pitcher plant (Sarracenia) as a modelsystem to examine three ecological themes. First, Jimevaluated cost-benefit models of plant carnivory bycalculating the construction costs of plant structures.It is assumed that there is a benefit to plants con-structing carnivorous structures, such as pitcher-shaped leaves to trap insects, but data to comparecosts and benefits is insufficient. Comparisons will bemade for multiple species of carnivorous plants in theHarvard Forest greenhouses collected from a range ofenvironments across North America. A second com-ponent examines the stoichiometry of carnivorousplants in response to prey capture and other nutrientsources, e.g., root uptake, atmospheric deposition. Amultinutrient approach is necessary because varyingnitrogen inputs influence the nutrient ratios of plants.For example, acidic deposition adds nitrogen withoutthe associated phosphorus and potassium inputs andcould create nutrient imbalances relative to plantsthat receive increased insect inputs that provide mul-tiple nutrients. The third component is a meta-analysis of nitrogen movement through food websfrom studies using N-15 stable isotopes. This analysiswill advance our understanding of the utility of N-15 stable isotopes to trace the movement of nitrogenthrough food webs and identify trophic levels.

PATTERNS OF FOREST HARVESTING,HISTORICAL LAND-USE, AND INVASIVESPECIES IN MASSACHUSETTS

Rob McDonald, David Foster, and Glenn Motzkincompleted a study evaluating the influence of forestharvesting, past land-use, and site conditions on thedistribution and abundance of invasive plant speciesin Massachusetts. Based on two seasons of field data,they determined that invasive plant species are wide-spread in the woodlands of central and westernMassachusetts, but typically occur in low abundance.The most common invasive species preferentiallyoccur on relatively rich soils and sites formerlycleared for agriculture. Extensive wooded areas havefew invasive species whereas woodlands in highlydeveloped or fragmented landscapes typically sup-port more invasives. The occurrence and intensity offorest harvesting over the past twenty years was nota strong predictor of the occurrence of several com-mon invasive species. Methodologies and resultsfrom this study will advance conservation planningacross the eastern United States, and will have broadrelevance for forest management and policy efforts.

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A page from the life story of nineteenth-century amateurbotanist Rebecca Merritt Austin (1832–1919), as told byher daughter Mary Hail to Frank Morton Jones in 1918.

Manuscript from the Frank Morton Jones Archives,Entomology Division, Peabody Museum of Natural

History, Yale University.

The project evaluating the broad ecological, con-servation, and resource implications of forest har-vesting across Massachusetts, initiated by GlennMotzkin, David Foster, and Dave Kittredge, hasachieved its major objectives. Previous work byAndrew Finley and John Burk had assembled a spa-tially explicit database of all forest harvesting in thestate since 1984.

Robert McDonald took the lead in quantifyingthe spatial pattern of forest harvesting in Massachu-setts. Mean harvesting intensity varies from 28.3 m3

ha-1 in the Taconics to 50.3 m3 ha-1 in the coastal plain,

with higher harvesting intensities generally occurringin the eastern portion of the state. Around 0.5 percentof forests in the coastal plain are logged each year,while 1.4 percent of forests are logged each year inthe Worcester Plateau. There are two main factorslimiting forest harvesting in the eastern portion of thestate: smaller parcel size, due to greater landscapefragmentation, and higher land prices, which reducesthe incentives to landowners to harvest their forestand increases the incentives to landowners to devel-op their land. This work was published in Forest Ecol-ogy and Manuscript in May 2006.

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Throughout Massachusetts, 24.1 percent of forests with a low road density (less than 2.5 km of road per km2)and a low house price (below $162,000 dollars) were harvested between 1984 and 2003. In areas of

the state where either of these two variables are greater, forest harvesting is much less prevalent.

Sites that were cleared for agriculture in the nineteenth century are twice as likely to have an invasive species present.

DISTRIBUTION, POPULATION DYNAMICS, AND IMPACTS OF INVASIVE PLANT SPECIES

Population-to-landscape level observations, historicalstudies, and controlled experiments are being con-ducted to investigate invasive plants as both respon-ders to and drivers of forest change. Although inva-sive plant species are widespread, remarkably little isknown about their ecological impacts, the landscapeto regional patterns of distribution and abundance ofthese taxa, or the factors controlling their rates andpatterns of spread. Recent results provide new evi-dence for direct and indirect impacts of the invasiveplant, garlic mustard, on native plants and their fun-gal mutualists. Geographic variation has also beendemonstrated in the distribution and abundance ofthis plant across ecoregions in Massachusetts, andlocal variation in the demographic performance ofinvading populations in different forest habitats.

The enemy of my friend: An invasive plant dis-rupts mutualistic associations between native treeseedlings and mycorrhizal fungi.

Very few studies have empirically investigated howinvading plants may alter ecological interactionsamong resident species in the invaded range. KristinaStinson and collaborators John Klironomos (Univer-sity of Guelph) and Ben Wolfe (OEB) demonstratedthat Alliaria petiolata (garlic mustard), a Europeaninvader of North American forests, suppresses the growth of key native canopy tree seedlings by disrupting their longstanding mutualistic associationswith arbuscular mycorrhizal fungi (AMF). Seedlingsof the dominant native hardwood tree species ofnortheastern North America (sugar maple, redmaple, and white ash) had lower growth and fungalcolonization of roots in soil invaded by garlic mustard than in uninvaded soils. These reductionswere similar to those observed on sterilized soils. Allthree species also demonstrated lower root coloniza-tion by AMF in soils conditioned by garlic mustardthan in unconditioned soils or soils conditioned byother native plants. The suppressing effects of garlicmustard appear to result from water-soluble chemi-cals exuded from the plant, as aqueous extracts fromgarlic mustard were just as effective as the livingplants at reducing the root colonization and germina-tion of AMF spores. Garlic mustard also suppressed

AMF colonization of roots in several additionalspecies typical of forest successional gradients. How-ever, tree seedlings were most strongly dependent onthe mutualism and demonstrated the strongest negative growth responses. Thus, the antifungaleffect of this invader disproportionately reduces theperformance of native tree seedlings over other plantspecies.

Landscape and habitat variation in demographicattributes of invasive plant species

Kristina, Kathleen Donohue, and summer under-graduates researched the role of landscape and habi-tat factors on demographic attributes of the invasiveplant Alliaria petiolata (garlic mustard) in northeasternforests. The Prospect Hill tract at Harvard Forest wassurveyed and they found that six invasive species,including garlic mustard, are most abundant in siteswith agricultural land-use history. Also, local popula-tions of garlic mustard were monitored in sun, inter-mediate, and forest habitats at Harvard Forest over aperiod of three years. Using census data from fourlife-stages in 90 1m2 quadrants, they constructeddemographic population matrices and found thatpopulation growth rate was greatest in sun popula-tions and lowest in intermediate habitats.

Survivorship between life-stages depended main-ly on yearly fluctuation and less on type of habitat.The reasons for the higher growth rate of sun popu-lations were higher seed production and a higher sur-vivorship from first- to second-year plants.

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Left: A Microscope view of a plant root colonized byarbuscular mycorrhizal fungi (AMF) spores; right:crushed spore of an AMF in the genus Acaulaspora.

Regional variation in forest invasion by garlic mustard

At a broader scale, Kristina, Brian DeGasperis, andREU student Kevin Burls recently conducted aregional comparison of landscape-level distributionof garlic mustard (Alliaria petiolata) in forested sites inMassachusetts. Using the Massachusetts Geographi-cal Information System (MassGIS) layers to deter-mine the locations of forest cover and subsequentlygenerate random sampling points, they surveyed fiftysites in two contrasting ecoregions, the ConnecticutRiver Valley and the Berkshire Valley for the follow-ing: presence/absence of garlic mustard in forest edgeand forest understory habitats, incursion rates (dis-tance of population invasion into forested habitat),and percent cover of this species. The Berkshire Val-ley had ~ 60 percent occurrence of garlic mustard,compared to only ~15 percent in the ConnecticutRiver Valley. Preliminary results demonstrate that

the two ecoregions differed in the percent cover offirst-year plants and in total percent cover of garlicmustard. Further analyses are underway to deter-mine the degree to which these locations differ inabundance and incursion rates, and the relative con-tributions of environmental, historical, and physio-graphic variation to the success of forest invasion.

Influences of historical land use and forest har-vesting on Japanese barberry invasion

Attempts to determine the characteristics of habitatsthat render them invasible to non-native species havemet with limited success. One possible explanationfor this shortcoming may be that by focusing onmodern conditions, most studies fail to consider inva-sibility in the context of a historically dynamic land-scape in which both the abundance of a species andthe invasibility of a site may change. Despite the rec-ognized importance of historical factors in controlling

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A forest understory in Massachusetts heavily invaded by garlic mustard.

many native species distributions, few studies haveincorporated historical landscape changes into models of invasive species distribution and abun-dance. As part of his MFS research Brian DeGasperisexplored the possibility that the current distributionof invasive species may reflect legacies of historicalland use despite nearly a century of forest successionand subsequent disturbances. This work is based inpart on the recognition that since the mid-nineteenthcentury southern New England has been trans-formed from a predominately agricultural landscapeto a largely forested condition. Thus, species in ourforests today may have spread and become estab-lished in fields or young forests of the past. Forestswere surveyed for the occurrence and abundance ofJapanese barberry, Berberis thunbergii, a problematicnonnative shrub in northeastern forests, relative totwo distinct periods of historical land use (1830 and1927), forest harvesting (1984–present), and environ-

mental and edaphic characteristics. Barberryoccurred more frequently and was more abundant insites historically cleared for agriculture than in con-tinually wooded sites.

This relationship was strongest for areas in agri-culture in the early twentieth century when barberrywas widespread in the region. While land-use historyis confounded with soil fertility and distance to puta-tive seed sources, Brian’s data suggest that the strongrelationship between modern distribution and priorland use is a consequence of colonization of aban-doned agricultural lands and persistence throughsubsequent reforestation. In contrast, recent forestdisturbances, such as harvesting, did not influencethe occurrence or abundance of barberry at the land-scape scale. These results indicate that invasibility ofa site must be considered in the context of changinglandscape conditions. In particular, interpretations ofboth native community composition and modern

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Legacies of an agricultural past: a wood land stonewall delineating a dense thicket of Japanese barberry.

plant invasions must consider the importance of his-torical landscape changes and the timing of speciesintroduction along with current environmental andedaphic conditions.

CO2 ENRICHMENT ALTERS REPRODUCTION, COMPETITIVE INTERACTIONS, AND GENETIC

Structure of Common Ragweed Populations

Plants growing in dense stands may not equallyacquire or utilize carbon gains from elevated CO2. Asa result, the size and reproductive differences betweendominant and subordinate plants may be alteredunder rising CO2 conditions. Kristina Stinson is com-pleting studies on the impacts of global change on thecompetitive interactions and relative reproductiveoutput of individuals and genotypes of the allergenicweed, Ambrosia artemisiifolia (common ragweed).

Effects of high CO2 on reproductive dominance

It is hypothesized that shaded, subordinate indivi-duals of ragweed would achieve greater growth andreproduction in proportion to stand totals than light-saturated dominants under elevated CO2. They grewexperimental stands of ragweed at either 360 µL L-1

or 720 µL L-1 CO2 levels and measured the growthand reproductive responses of competing individuals.To test whether elevated CO2 diminished or exag-gerated size and reproductive inequalities withinstands, stand-level coefficients of variation (CV) werecompared in final shoot, root, and reproductiveorgan biomasses. Smaller plants produced morereproductive biomass per unit of growth in theenriched-CO2 treatment. The elevated CO2 treat-ment stimulated growth and reduced inequalities inall plant organs, especially reproductive allocation.Positive relationships between the CV and total standproductivity deteriorated under elevated CO2, indi-cating that growth enhancements to smaller plantsdiminished the relative biomass advantages to largerplants in increasingly crowded conditions. It was con-cluded that CO2-induced growth gains of subordi-nate A. artemisiifolia plants stimulate stand-level repro-duction and equalize the reproductive output of smalland large plants. Thus, more individuals are likely toproduce greater amounts of seeds and pollen infuture populations of this allergenic weed.

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Frequency of occurrence of Japanese barberry in modernforests relative to: (a) land use prior to barberry’s intro-

duction to Massachusetts; (b) post-introduction land use;and (c) modern harvesting activity. The distribution ofbarberry is strongly influenced by historical agriculture,

but not by recent selective logging.

Architectural and physiological response to high CO2

Using data from the experiment described above,REU students Jen Petzold and Jimmy Tran com-pared the relative growth, photosynthetic capacity,and architecture of subordinate and dominant plantsin each treatment, and assessed size inequalities usingthe stand-level coefficient of variation (CV). In ele-vated CO2, plants grew larger, but subordinate plantsshifted more mass to upper stem allocation thandominants. Dominant plants demonstrated reducedleaf-level photosynthetic gains in elevated CO2 com-pared to subordinate plants. Reduced CVs in plantsize reflected smaller proportional growth gains bydominants over subordinates in elevated versusambient stands. Jen and Jimmy concluded that dif-ferences in the architectural and physiologicalresponses of subordinate and dominant ragweedplants reduce competition and allow subordinateplants to catch up to dominants in elevated CO2

conditions.Functional relationship models for the biomass

and reproductive allocation of ragweed genotypes atambient and elevated CO2.

Using functional relationship models, Kristinaand collaborators at University College, Dublin, Ire-land, John Connolly and Caroline Brophy, are inves-tigating the effect of CO2 on the hierarchical structureof genotypes within these stands with respect to bio-mass and reproductive allocation. Twelve genotypesof ragweed from known maternal lines were selectedand grown competitively in the experimental standsdescribed above. Each genotype was replicated twice

within each stand. To date, they have found novelevidence that the structure of populations changedunder elevated CO2 — those genotypes that performless well under ambient CO2 conditions benefit pro-portionately more than others under elevated CO2,i.e., subordinate genotypes ‘catch-up’ to dominants.This was true for both biomass and reproductiveallocation. Moreover, genotypes with a smaller repro-ductive allocation at ambient CO2 were no longersubordinate under elevated CO2 as all genotypestended to a common reproductive allocation. This isthe first study to demonstrate a genetic basis for dom-inance hierarchies within developing stands, and toshow how elevated CO2 can alter these hierarchies toaffect the outcome of competition.

HYDROLOGICAL STUDIES

The combination of long-term hydrological, meteor-ological, and eddy flux measurements at the HarvardForest provides a rare opportunity to study thehydrological cycle of small forested watersheds. Apilot project conducted by Emery Boose, JulianHadley, and summer undergraduate Safina Singh insummer 2005, estimated water budgets for ProspectHill by integrating precipitation data from the Fishermeteorological station, evapotranspiration data fromthe Hemlock and Little Prospect Hill eddy flux tow-ers, stream discharge data from the Nelson andBigelow gages, and ground water data from wellsinstalled in the 1970s by Walter Lyford. In bothwatersheds results showed the expected seasonaldecline in system water storage from late springthrough early summer in both the Nelson and UpperBigelow watersheds, as well as a significant increasein the relative loss of water to evapotranspirationafter leaf-out in early June. Nelson watershed, with itsmuch higher percentage of wetland area (60 percentvs. 10 percent), retained significantly more water dur-ing this period. In both watersheds the change inwater storage calculated from precipitation, evapo-transpiration, and stream discharge closely matchedthe estimated change in ground water storage.

Recent developments in wireless networking andsensor technology will make it possible to collect,analyze, and display such measurements online innear-real time. However management of such datastreams presents a number of interesting technicalchallenges, including how to do real-time documen-tation and quality control of the data. These prob-

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Culvert built in the fall of 2006 on Bigelow Brook.

lems are being studied as part of a long-term collabo-ration between ecologists at the Harvard Forest:Aaron Ellison, Emery Boose, and Julian Hadley, andcomputer scientists at UMass-Amherst, Leon Oster-weil, Lori Ann Clarke, Alexander Wise, using a real-time water budget system for Prospect Hill as amodel case.

ECOLOGY AND CONSERVATION OFKETTLE PONDS AND VERNAL POOLS

Efforts to complete long-term studies of vernal poolsin Massachusetts have continued. Betsy Colburnanalyzed five years of data from fourteen vernalpools on Cape Cod and prepared a manuscriptexamining patterns in community change in relationto year-to-year variations in precipitation and poolhydrology. Betsy also continued processing of sam-ples collected from vernal pools across the state, as

part of an ongoing comprehensive analysis of inver-tebrate community composition in relation to geog-raphy, hydrology, and historic land use.

As part of the development of long-term moni-toring methods for kettle ponds and vernal pools inthe Cape Cod National Seashore, Betsy carried out a

multivariate comparison of kettle ponds in relation totheir chemistry, basin characteristics, and other limnological variables. Processing of aquatic inverte-brate samples from the ponds is continuing, and theresults will ultimately be combined with the physical-chemical data.

STREAM ECOLOGY AND POTENTIALIMPLICATIONS OF HEMLOCK DECLINEMEDIATED BY THE HEMLOCK WOOLLYADELGID

Harvard Forest Aquatic Ecologist Betsy Colburn andBill Sobczak from College of the Holy Cross’s Biolo-gy department continued baseline research on head-water streams in north central Massachusetts, in con-junction with ongoing forest studies by Dave Orwigand other Harvard Forest researchers. Former 2004REU student Bridget Collins worked with Bill to pre-pare a manuscript for submission to Wetlands, detail-ing the results of her examination of chemical andbiological characteristics of surface reaches and sub-surface flow chutes in Bigelow Brook.

In summer 2005, Betsy Colburn and Bill Sobczakco-mentored two summer REU students: JamesWillaker from the SUNY College of the Environmentand Forestry and Matthew Kaufman from Keene StateCollege. The students carried out intensive habitatsurveys in headwater streams in northern Worcestercounty, adding substantially to our database on phys-ical, hydrological, and watershed characteristics ofstreams in hemlock-dominated and deciduous water-sheds. In addition to collecting standardized streamhabitat data, James and Matt applied standard forestry

The big weir on Nelson Brook.

13

Emery Boose on a new weir.

14

Boose, Hadley, and Singh. Preliminary Water Budget for Prospect Hill

Late Spring 2005

P P

P P

ET ET

ET ET

Mill

imet

ers

Mill

imet

ers

Q

QQ

Q

dS

dSdS

dSdGW

dGWdGW

dGW

Nelson

Nelson

Bigelow U

Bigelow U

Early Summer 2005

6

0

-6

6

0

-6

p

Preliminary water budgets for Nelson and Upper Bigelow watersheds for late spring (April 20–June 5) and earlysummer ( June 6–July 26) 2005. Values are daily averages. Positive values represent a gain and negative values a loss ofwater to the forest ecosystem. P = precipitation measured at Fisher Meteorological Station. ET = evapotranspirationmeasured at Hemock and Little Prospect Hill eddy flux towers, with values weighted according to the proportion ofhemlock and hardwoods in each watershed. Q = yield measured at steam gages, with gaps in data filled by simple runoffmodel. dS = chnge in ecosystem wter storage, calculated as dS = P + ET + Q. dGW = change in ground water stor-age measured at two nearby wells chosen to represent poorly-drained and well-drained soils, with values weightedaccording to the percentage of wet and dry soils in each watershed. (Boose, Hadley, and Singh, unpublished data.)

survey techniques to quantify forest composition inthe catchments of study streams. This information,along with results from previous and ongoing surveys,serves as a solid baseline for future comparisons if andwhen the hemlock woolly adelgid (HWA) becomesestablished in northern Massachusetts and kills hem-locks along headwater streams. Dr. Sobczak contin-ued much of the survey work through the year withstudents in his undergraduate aquatic ecology class.

As an independent project, James Willaker com-pared the aquatic invertebrate communities in theeastern and western branches of Bigelow Brook,respectively deciduous and hemlock-dominatedheadwater streams at the HF Prospect Hill Tract, andcontinued his analysis of the communities as a seniorthesis during the following year. For his project, MattKaufman installed two parallel transects of ground-water wells perpendicular to the upper channel of thewest tributary of Bigelow Brook, and examined rela-tionships between streamflow, groundwater eleva-tions, and precipitation. The wells remain in situ forfuture research in this area.

Betsy compared Dave Orwig’s digitized aerialphotographic data on the distribution and density ofhemlock and hemlock woolly adelgid in Massachu-setts with topographic map evidence of stream chan-nels, and identified a number of potential candidatestreams for focusing future research on short-termimpacts of HWA on streams. In summer 2006, fieldsurveys by Bill Sobczak and two REU students, LoriSaunders and Tim Rowell, are focusing on streamsdominated by hemlock and affected by HWA.

In spring, 2006, Betsy quantified light inputs todeciduous and hemlock streams and, with JoanMilam, sampled mosses from the eastern and west-ern branches of Bigelow Brook. The results, alongwith an overview of habitat and community datafrom Harvard Forest headwater stream surveys todate, were presented in a poster at the North Ameri-can Benthological Society Annual Meeting inAnchorage, Alaska, in June 2006.

CHANGING DYNAMICS OF HEMLOCKWOOLLY ADELGID POPULATIONS AND ASSOCIATED IMPACTS ACROSSSOUTHERN NEW ENGLAND

As part of a larger Harvard Forest research emphasison invasive species, work continued on several stud-ies led by Dave Orwig investigating the impact of theintroduced insect pest, the hemlock woolly adelgid.

During the summer of 2006, Dave and Heidi Luxcontinued to examine the response of ecosystemprocesses to the stress and mortality caused by theHWA in Connecticut hemlocks. Over the eight-yearstudy differences in nitrogen availability betweenheavily infested and lightly infested sites were obvi-ous for the first four years then recently declined,probably due to HWA declines resulting from coldwinter temperatures November 2004 through spring2005, and November 2003 through spring 2004.Interestingly, overstory mortality has continued atsites in central and northern Connecticut (15 percentto 35 percent in 2005), but not at the rates experi-enced in southern Connecticut (all sampled sites havenow lost >80 percent overstory hemlock). Daveworked with summer student Christie Rollinson(Oberlin College) to establish additional permanentplots in Massachusetts to follow the long-term forestdynamics associated with HWA infestation at thenorthern edge of the invasion.

As a follow-up to examining the distribution ofhemlock, HWA, and HWA-related damage acrossthe landscape of Connecticut and Massachusetts, alarge resampling effort has now been completedacross our 7,500 km2 study area. We assessed thespread and community-level impact of the HWA andanother invasive insect attacking hemlock, the elon-gate hemlock scale (EHS), on southern New Eng-land hemlock forests by resurveying 141 hemlockstands. All of these stands had been previously iden-tified via aerial photography and surveyed in either1997–98 (CT) or 2002–04 (MA) for HWA and EHSdensity as well as hemlock stand vigor. We rated twobranches on each of fifty trees per stand on a scale of0–3 (0=none; 1=1–10 organisms/m branch;2=11–100/m branch; 3=100+/m branch) for HWAand EHS density. We also assessed percent overstoryhemlock mortality as well as percent canopy loss ineach stand.

Canopy loss and hemlock overstory mortalityincreased with increasing HWA density, with themost damaged stands located in southern Connecti-cut and the most undamaged stands in northernMassachusetts. HWA density decreased with increas-ing latitude, possibly due to climatic limitations, andwas absent from much of northern Massachusetts.Although the total number of HWA-infested sitesincreased between the initial and 2005 surveys, meanHWA density per site has decreased over this sameperiod, perhaps due to extreme cold spells in 2004and 2005. EHS distribution and density per site has

15

increased dramatically since 1997–98 and is nowpresent in every Connecticut stand and many Mass-achusetts stands. Within the survey transect, EHSappears to be moving rapidly northeast from the siteof its initial introduction on Long Island in 1908.Despite its rapid increase in abundance, EHS densi-ty was not correlated with overstory hemlock mor-tality when the effect of HWA presence was takeninto account. There was a positive correlationbetween HWA and EHS density on trees within indi-vidual sites in sites with low overall HWA densities.As overall HWA density increased, however, this cor-relation shifted from positive to negative (likely as aresult of resource competition). This large project is acollaborative effort that includes Dave, undergradu-ate student Sasha Lodge, Kenyon College; EvanPreisser, University of Rhode Island; and Joe Elkin-ton, University of Massachusetts–Amherst.

Forests in Massachusetts and Connecticut areincreasingly affected by the invasion of the hemlockwoolly adelgid (Adelges tsugae) both through directeffects of mortality from infestation and from salvagelogging. Over the past five years the impact of sal-vage logging in these systems has been studied byour group and, in some cases, it has been clear thatlogging of damaged stands has a greater impact onecosystem function than the effects of deteriorationdue to adelgid driven mortality alone.

For example, an infested site that was cut in 2002has been replaced with black birch, raspberries, andother herbaceous species. Soil temperature and nitro-gen availability have increased tremendously sincethe cutting and exceed the values observed in any ofthe infested forests. Organic soil moisture at the cutsite has continued to decrease since the logging. Wewill monitor these sites with ion-exchange resin bags

to examine the spatial availability of N within sitesand the extent to which NO3 is being lost over time.

In 2003, Heidi Lux and Dave Orwig began tostudy post-adelgid logging and ecosystem function atthe urban Arnold Arboretum, just outside of Boston.During 2004, six 15 × 15 m plots were fenced off,and baseline data on soil nutrient cycling, microcli-mate, and vegetation information were collected.Hemlocks were removed from four plots in Februaryof 2005, while two remain untouched as controlplots. Slash was chipped and left on site in thechipped treatment plots or removed from the logged(only) treatment. The logging operation was con-ducted by crane and had low impact on the soils.This series of treatments coincides with those at theHarvard Forest’s Hemlock Removal Experiment atthe Simes Tract in Petersham, where they are collect-ing analogous data.

Dave and Heidi are currently in their secondyear of post-treatment measurements at both of thesesites, following one year of baseline data collection.In general, both nitrogen availability and nitrogencycling rates are very high at the Arboretum, partic-ularly for nitrate. This was true even before treat-ments began, with availability and rates from two totwenty times higher than had been observed at themore rural study sites. Nutrient cycling responses arelikely to be even more dramatic this growing seasonand next, based on observations at logged, infestedsites in Connecticut, where the mobilization of nutri-ents peaks in the second and third years followingcutting. Vegetation responses to cutting at theArboretum have been rapid and dramatic, withshoulder-high cover of up to 50 percent in the sum-mer of 2005, just months after cutting. While theyexpected a proliferation of weedy and shrubby

16

Heidi Lux at the Arnold Arboretum in 2004. Heidi in the same area in 2005!

species, Dave and Heidi were astonished by the dra-matic response as vegetation at other logged sites hastaken a year or two to establish significantly.

Soil nutrient cycling and vegetation responses atthe Harvard Forest Simes tract were muted in the firstyear post-treatment, and in line with other commer-cially logged sites. The next several years will presentmany opportunities for comparison between theurban and rural study sites.

Many landowners are choosing to pre-emptivelyharvest their hemlock stands with the advent orthreat of HWA. Information from timber harvesters,state agencies, and Harvard Forest studies indicatethat the recent broadscale increase in logging associ-ated with HWA is occurring with little ecologicalassessment and in the absence of scientific back-ground for conservationists, land managers, or policymakers. Dave Orwig and David Kittredge haveincorporated findings from several previous andongoing Harvard Forest studies of HWA and hem-lock logging into a pamphlet for landowners that pro-vides various silvicultural options for managing hem-lock forests. Options range from doing nothing tolight selective cutting to high-intensity cutting,depending on landowner objectives, overall hemlockhealth, and stand conditions. In addition, the pam-phlet includes planting options and addresses HWAbest-management practices. All options and costsshould be considered carefully when planning theappropriate management strategies to effectivelymeet desired goals and make an informed decisionabout hemlock and the threat of HWA.

Local high-school graduate Eowyn Connelly-Brown assisted for a second year with many of thelaboratory and field activities associated with theselabor-intensive studies. In addition, Shelley Ray-mond, an undergraduate of the University of Massa-

chusetts–Amherst, volunteered during the summerof 2006 on several HWA-related projects.

EXPERIMENTAL LOGGING ANDGIRDLING OF HEMLOCK

To complement the extensive work on the HWA andexperimental plots at the Arnold Arboretum, in2003, Aaron Ellison, Audrey Barker Plotkin, andDave Orwig began a large-scale Hemlock RemovalExperiment to simulate the effects of hemlockremoval from the forest ecosystem by adelgid and bylogging. There are two replicates (0.8 ha plots) of thefollowing treatments: hemlock girdling (to simulatemany of the effects of HWA), hemlock commerciallogging (to simulate the preemptive logging that isoccurring in hemlock forests in our region), hemlockcontrol (no treatment), and hardwood control (representing a possible future composition of post-hemlock forest). Logging and girdling treatmentswere completed in winter and spring 2005.

The Hemlock Removal Experiment is now fullyup and running. Ongoing measurements includehourly measurements of soil and air temperature (byAaron Ellison — HF); twice-yearly measurements oflight quantity from hemispherical canopy photo-

17

Dave Kittredge and REU student Kelly Grogan.

Available nitrate at the experimental plots on HemlockHill, at the Arnold Aboretum (a) and at the Hemlock

Removal Experiment at the Simes Tract, Harvard Forest(b). The first bar on the left of each group in b. represent

an additional collection of pretreatment data in the winter of 2003–2004. Manipulations began at

both sides of the winter-spring of 2005.

graphs (Aaron Ellison); soil respiration (KathleenSavage, Emily Austin, and Eric Davidson — WHRCand Hampshire College); nitrogen availability andmineralization rates (Heidi Lux and Dave Orwig —HF); throughfall chemistry (Pamela Templer — BU);primary productivity (litterfall) and rates of decay ofcoarse woody debris (Audrey Barker Plotkin — HF);species composition of understory herbs and shrubs(Audrey Barker Plotkin); ants and beetles (AaronEllison, REU students Laura Briscoe, Grace Wu,Alex Arguello, and Rachel Osborn, and UMassPh.D. student Sydne Record); birds (volunteers Ernieand Linda LeBlanc of Royalston), and salamanders(Brooks Mathewson — HF); overstory tree size andvigor (Audrey Barker Plotkin). Data are being usedto parameterize models forecasting changes in netecosystem productivity in response to adelgid infes-tation (Paul Moorcroft — OEB). Parallel experimentsare underway at the Coweeta LTER site (hemlockremoval) and the Black Rock Forest in eastern NewYork (oak removal).

Initial data, including two-year-before treatmentsand one-year-after treatments, show clear changes inmicroclimate, plant and animal species composition,and ecosystem properties in logged plots vs. controls.Girdled hemlocks began to disintegrate in summer2006, and although girdled plots are noticeablybrighter due to thinning of the hemlock canopy andfalling branches and trees, few measured variablesshow significant responses to treatments.

Eric Davidson and Kathleen Savage (WHRC)are examining the change in carbon released thor-ough soil respiration as a result of hemlock death.The balance between the components of soil respira-tion (root respiration and microbial respiration) mayshift during the disturbance period. With the girdling

treatment they can see how soil respiration rateschange as the hemlocks die. This will also giveinsight into how to partition between the root versusmicrobial components of soil respiration; microbialrespiration should dominate in girdled plots. In col-laboration with Sue Trumbore from the University ofCalifornia, Davidson’s group will also look at theC14 signatures from the girdled experiment andcompare those to the control stand, which will helpdetermine the age of the carbon being respired.

Aaron presented preliminary result of this largecollaborative experiment in a poster at the triennialLTER All Scientist’s Meeting in September 2006,and in a workshop at the meeting on foundationspecies that he organized with Brian Kloeppel fromCoweeta LTER.

Ant controls on ecosystem dynamics

With a lot of backbreaking labor contributed by JessButler, Jim Karagatzides, Emily Austin, and TonyD’Amato, Aaron also established a set of 3 × 3-m antexclosures and disturbance controls to determine ifants really are the “little things that run the world” orat least run the soil ecosystem dynamics at the SimesTract. This experiment builds on observations pub-lished by Walter Lyford in 1963 (Harvard ForestPaper no. 7) and is also being replicated at Coweetaby Aimée Classen (Oak Ridge) and Nate Sanders(University of Tennessee).

Doctoral student Sydne Record and 2006 REUstudent Alex Arguello are working with Aaron tocompletely inventory the ant diversity of the SimesTract. Focal areas for sampling in summer 2006 werebased on unique vegetation identified by Jerry Jenk-ins and Glenn Motzkin in their Flora of Harvard For-est project. 2006 REU student Rachel Osbornworked with Aaron to extend work by 2005 REUstudents Grace Wu and Laura Briscoe to inventorykey groups of beetles — carabids, cerambycids, andelatyrids — at the Simes Tract. In the long run, thiswork will contribute to an inventory of the insectfauna of Harvard Forest.

FOREST EXCHANGES AND INTER-ACTIONS WITH THE ATMOSPHERE

New heating x warming experiment

Serita Frey from the University of New Hampshirereceived a NSF Career grant to establish a new

18

REU students Dave Diaz and Chelsea Kammerer-Burnham with Aaron Ellison.

experiment at Harvard Forest to examine the inter-active effects of soil warming and N additions onmicrobial community structure and nutrient cyclingdynamics. The plots were established in fall 2005.There are four treatments (control, heated +N, heated −N, +N only) and six replicates in a com-pletely randomized design (twenty-four 3 × 3 mplots). Average soil temperature in the heated plots iselevated 5º C above ambient by the use of buriedheating cables placed at 10 cm depth in the soil andspaced 20 cm apart. The heating cables are con-trolled by a data logger that monitors thermistors ineach plot every ten minutes. Plots automatically turnon and off to maintain a 5º C temperature differencebetween the heated and control plots. The N addi-tion plots (heated only, +N only) are fertilized fol-lowing the protocol of the Chronic N Addition Study.An aqueous solution of NH4NO3 is applied at a rateequivalent to the low N plots at the chronic N study(5 g m-2 yr-1). Fertilizer is being applied in equalmonthly doses during the growing season (Aprilthrough October). The control plots and unfertilized,heated plots (heated only) receive water only. Rou-tine measurements will include soil respiration, Nmineralization, microbial biomass, and metabolism,and C and N pools (total C and N, dissolved organ-

ic C, inorganic N). This study will experimentallyshow how anthropogenic change, i.e., warming, Ndeposition affects microbial community compositionand diversity, and how shifts in the microbial com-munity impact C and N cycling at the ecosystemlevel. Serita is developing a new approach(calorespirometry) to assess microbial metabolism,with an emphasis on carbon-utilization efficiency.Alix Contosta, a doctoral student in Serita’s lab, willfocus her dissertation on the fungal community andwhether shifts in fungal species composition, inresponse to warming and/or N additions, are linkedto changes in litter decomposition rates.

Soil respiration studies

In 2005, Eric Davidson and Kathleen Savage fromthe Woods Hole Research Center began theireleventh year measuring soil respiration at the Forest,and added two new plots to their long-term transectsites. These long-term studies have demonstratedlarge differences among years in fluxes of CO2 fromthe soil. The fluxes tend to be much lower duringyears with summer droughts and higher during wetsummers. Recent expansion of measurements inareas to the north and east of the EMS tower also

19

Ant exclosure.

show important spatial heterogeneity, with lowerfluxes from the shallow soils on steep slopes in theseareas. They are working toward an understanding ofhow to integrate the spatial heterogeneity of soil CO2

efflux with the tower-based measurements of totalecosystem respiration.

Fast time response measurements of volatileorganic compound (VOC) concentrations and fluxes at Harvard Forest

Karena McKinney, Assistant Professor of Atmos-pheric and Environmental Chemistry at AmherstCollege, continued measurements of volatile organiccompounds (VOCs) at the Harvard Forest Environ-mental Measurement Site. Through reactions withthe major atmospheric oxidizing agents and NOx,VOCs play a key role in determining the oxidizingcapacity of the atmosphere. The goal of this project isto quantify sources and sinks of VOCs at the Har-vard Forest and investigate their reactivity in theatmosphere by making direct field observations ofambient concentrations, concentration gradients, andfluxes of VOCs.

During the summer and fall of 2005, measure-ments of many VOCs, including isoprene, monoter-penes, methyl vinyl ketone, methacrolein, benzene,tolulene, xylenes, acetone, and acetaldehyde wereobtained. Biogenic species such as isoprene andmonoterpenes are the dominant VOCs over HarvardForest in the summer, although measurable quantitiesof anthropogenic species are also present, indicatingthat the forest atmosphere is impacted by anthro-pogenic hydrocarbon emissions from nearby urbanand industrial areas. Diurnal cycles of isoprene andits oxidation products, methyl vinyl ketone andmethacrolein are being used to understand the fate offorest emissions, which are both removed chemicallyand redistributed through dynamical processes. Therelative importance of these mechanisms affects theproduction of ozone due to VOC oxidation in thevicinity of the forest. Additional measurements ofVOC concentrations, concentration gradients, andfluxes, took place in 2006.

NATIONAL INSTITUTE FOR GLOBALENVIRONMENTAL CHANGE

Julian Hadley continued to measure forest-atmos-phere exchange of carbon and water at the LittleProspect Hill and Hemlock forest flux measurement

towers, and completed analysis of data through theend of 2004. In the deciduous forest at Little ProspectHill, which largely established after a 1957 forest fire,higher annual carbon storage was observed in 2004compared to either of the preceding two years. How-ever, annual estimated net C storage at LittleProspect Hill was again similar to the value calcu-lated for the Environmental Measurement Site(EMS) tower, which is surrounded by older forestencompassing a wetland area and moister soils. In2004 as in 2003, both the maximum rate of carbonstorage above ground and the average rate of carbonproduction in the forest (ecosystem respiration) werelower for the younger, drier forest on Little ProspectHill compared to the EMS forest. These two differ-ences again roughly balanced each other, resulting insimilar net carbon storage. Carbon uptake by thehemlock forest in the summer of 2004 was muchlower than of the deciduous forests, and was similarto past measurements.

In the summer of 2005, Julian worked with twosummer students, Susan Cheng and Jennifer Mc-Innis, to measure aboveground carbon storage onLittle Prospect Hill using tree-ring analysis and vari-ation in dissolved carbon dioxide content of theBeaver Swamp northwest of the EMS tower. Thetree-ring analysis showed that C storage increasedrapidly following the 1957 forest fire as a new forestwas established, but that there was a sharp drop instorage in 1980 and 1981, probably due to severedefoliation by gypsy moths. The study of dissolvedCO2 levels in the Beaver Swamp was stimulated byunusually high rates of CO2 release observed inwinds blowing over the Beaver Swamp. Data fromthe swamp in 2004 showed large fluctuations in dis-solved CO2 concentration, ranging from above15,000 parts per million (ppm) to less than 4,000ppm. An experiment using a floating wind tunnelalso showed that the flux of CO2 from swamp waterto the atmosphere increased sharply as wind speedabove the water’s surface increased. This pattern isconsistent with the observations of large carbonreleases from the direction of the swamp.

STUDIES IN TREE AND PLANT BIOLOGY

Biophysics of plant form and function

Research conducted collaboratively by the N.Michele Holbrook, Department of Organismic andEvolutionary Biology and Maciej Zwieniecki, Arnold

20

Arboretum, focuses on the biophysics of plant formand function with an emphasis on the vascular sys-tem of trees.

Research over the past year, led by post-doctoralfellow Brendan Choat, addressed the role of ion-mediated changes in xylem hydraulic conductivityon water transport in sugar maple trees. Assisted byREU student Melissa Whittaker, this projectexplores the hypothesis that potassium ion (K+)concentrations in the xylem are up-regulated inbranches experiencing good condition for photosyn-thesis, possibly as a result of recirculation from the

phloem. The group recorded significant differencesin K+ concentrations in relation to both canopy posi-tion and light environment, but the predicted impacton xylem hydraulic conductivity is small. They havealso recorded substantial variability in both the sen-sitivity by which different branches respond tocation concentrations — the significance of which isunknown — but which seriously complicates analysisof these interactions. Further work will focus onunderstanding the source of this variation in sensi-tivity and its possible connection with wound-relatedresponses of cut branches.

During summer 2006, REU student KayaSchmandt continued work on the xylem vulnerabil-ity of red oak leaves. Capitalizing on the exception-ally wet spring, Kaya has documented a significant

shift in the vulnerability of red oak leaves, comparedwith data collected in 2003. Maximum leaf hydraulicconductivity, however, was higher in the wetter year(2003). Such developmental plasticity is somewhatunexpected, and suggests that there might be signifi-cant costs associated with building leaves that aremore resistant to cavitation and/or tradeoffs betweenhydraulic safety and efficiency.

Physiology of water transport in trees all year round

Tree stem diameter varies diurnally because ofchanging water tension inside the stem, which is ameasure of water availability for transpiration.Sanna Sevanto, a post-doctoral researcher in MissyHolbrook’s lab, measured this variation on severalspecies at Harvard Forest in order to evaluate thediameter variation method against other methodsfor measuring sap flow and transpiration and to shedlight on physiology and tree-level dynamics of watertransport of those species. Diameter variations aremeasured using linear displacement transducersattached to tree stems. The measurements are accu-rate and easy to maintain and the two water path-ways in trees (xylem for water and phloem for sug-ars) can easily be detected separately, which makesthem an interesting new approach to detect tree-levelwater dynamics through all seasons.

Sanna’s work at Harvard Forest consisted ofthree projects: 1) Diameter variations vs. sap-flowmeasurements in collaboration with NathanPhillips’s group from Boston University, 2) Watertransport during leaf fall and budbreak in collabora-tion with Steve Wofsy’s group from Harvard Uni-versity and 3) Phloem-xylem water transportdynamics. All the projects included measurementson red maple, paper birch and black birch. In proj-ects one and two red oaks were also measured.

The results of these projects show that in well-watered conditions the daily amplitude of xylemdiameter variation per water lost is directly propor-tional to the content of living cells in xylem.

This indicates that the elasticity of the xylem tis-sue is determined by the amount of living cells andthat water tension is readily transmitted between thedead conduction vessels and the living cells, whichmeans high cell-vessel conductivity. In all the speciesphloem contributed about two thirds to the totaldiameter variation of the stem, which also can beexplained by the higher content of living cells of

21

View of EMS tower.

phloem tissue. Project three showed that the radialconductivity between xylem and phloem is fairlyhigh and the sugar transport modifies diurnalphloem diameter only slightly. This was supportedby the results from project three where in the absenceof transpiration, i.e., leaf-less trees, the phloem diam-eter was observed to swell during the day as a resultof translocation of sugars. In contrast, during sum-mertime with full transpiration phloem shrinks withthe xylem during daytime. The wintertime swellingof phloem results in either local or overall water cir-culation in the stem. In the absence of transpiration,local water circulation might be the only oxygen sup-ply for living cells in the xylem. However, maintain-ing overall circulation throughout the winter wouldserve the same function as well as provide means oftransporting sugars from stem reserves fuel to the liv-ing cells in the roots. Roots are the largest complex ofliving tissue in deciduous trees wintertime and wouldneed resources stored elsewhere to remain functionalthrough the winter.

LONG-TERM CHANGES IN CLIMATEAND VEGETATION

The paleoecology laboratory at the Harvard Forestexplores long-term changes in climate and vegetation

by analyzing sediment cores from lakes and ponds.Researchers and collaborators, including WyattOswald, Elaine Doughty, David Foster from the Harvard Forest, Jason McLachlan the University ofNotre Dame, and Barbara Hansen from the Univer-sity of Minnesota are currently synthesizing pollendata from across southern New England in an effortto reconstruct regional patterns of vegetation. In onestudy, comparison of several coastal and inlandrecords reveals contrasting dynamics in response toclimate change at ~5500 years ago. Similarly, in-depth analyses of records from central Massachusettsand the coastal region show spatial variations in veg-etation related to climate, disturbance, substrate, andphysiography.

Undergraduate researchers in the Harvard ForestREU program have made important contributions toour efforts this year. Sarah Truebe, student at Stan-ford University, analyzed a pollen record from Ben-son Pond in the Berkshires, and greatly improved ourunderstanding of the vegetation history of westernMassachusetts.

Alex Ireland, a student from Clarion Universityused tree-ring data, historical records, and a sedimentcore from a small pond to reconstruct the past

22

REU student Kay Schmandt dyeing leaves.

The diameter variation measurement system. The sen-sors (LVDT: Solartron Inc, UK) are attached to a rigidmetal frame mounted around the stem. For measuringxylem diameter variation (the right-hand side sensor)two screws are screwed through the bark on oppositesides of the stem. The sensor tip rests on one and theframe on the other. The sensor on the left measures

diameter variation on the bark (phloem included) andcomparisons of xylem and whole-stem diameter varia-tions reveal information of xylem-phloem interactions.

The cable is a thermo-couple measuring frame tempera-ture for the correction for thermal expansion of the

frame and the cover protects the system from heating bydirect sunlight.

23

changes in forest composition in the French Roadarea of Prospect Hill. His work provides an interdis-ciplinary view of post-agricultural forest dynamics ina little-studied part of the Harvard Forest.

Harvard Forest paleoecological research is alsoexpanding to other parts of New England. Sedimentcores recently collected from Umpawaug Pond andHighstead Swamp will provide long-term context for studies of the land-use history and present-day ecosystems in Redding, Connecticut, an oak-dominated region of New England that has beenignored by paleoecologists. Similarly, paleo lab mem-bers are collaborating with Kendra McLauchlan andJoe Craine of Dartmouth College on a study of pastecosystem processes in the White Mountains of NewHampshire. Geochemical analyses of a new sedimentcore from Mirror Lake are being used to reconstructchanges in nitrogen availability, a study that will com-plement decades of research by Gene Likens andHubbard Brook researchers on the lake ecosystem.

STRUCTURE, DYNAMICS, AND PROPERTIES OF OLD-GROWTH FOREST ECOSYSTEMS IN WESTERN MASSACHUSETTS

Tony D’Amato, a doctoral candidate working withDave Orwig, continued studies examining the com-position and structure, disturbance dynamics, andecosystem properties of old-growth forest systems inwestern Massachusetts. In these studies, eighteen old-growth and eight second-growth forest stands havebeen intensively sampled to determine the range ofvariation in ecosystem attributes, including coarsewoody debris (CWD) abundance, disturbancedynamics, and nitrogen cycling patterns among east-ern hemlock (Tsuga candensis) dominated old-growthand second-growth forest ecosystems.

This work has demonstrated that these old-growth forests contain a higher degree of structuralcomplexity than second-growth forests and that vari-ation in structural attributes, such as CWD abun-dance and tree size distributions, among old-growthstands is strongly correlated with disturbance history.In particular, old-growth forest stands experiencinghigher rates of natural disturbance over the past 130years, e.g., 5.4–9.9 percent canopy area disturbed perdecade, tend to have higher levels of structural com-plexity indicating the importance of natural distur-bances in generating many of the structural attributescommonly associated with old-growth forest ecosys-tems. Currently, information gained through thisresearch is being used to assist State planning efforts

Diurnal diameter variation on a maple tree a) with full leaves (September 18, 2005) and b) with no leaves (October 30, 2005). The phloem starts to swell during daytime when about half of the leaves are present.

Phloem curve is calculated by subtracting xylem from the measurement on the bark.

REU student Sarah Truebe and Wyatt Oswald.

24

for large-scale reserves on public land in Massachu-setts, as well providing a framework for efforts aimedat developing silvicultural strategies for restoring old-growth forest attributes to managed systems.

HISTORICAL ECOLOGY IN REGIONALSTUDIES

Drivers of mid-nineteenth-century woodlandlocation in coastal New England

Forests on the coastal islands of southern New En-gland and New York are important for the region’sbiodiversity due to their high number of unusualplant and animal species and communities. Ecolo-gists at Harvard Forest are gaining an appreciationof the importance that historic landscape patternsplay in shaping modern vegetation patterns.Therefore conservationists and land managersmust consider the land-use history of these foreststo explain current vegetation patterns. As part ofan effort to more fully understand the coastal land-scapes of Long Island, Block Island, Martha’sVineyard, Nantucket, and Cape Cod, Brian Hall,Glenn Motzkin, and David Foster have been using

the U.S. Coast and Geodetic Survey maps from themid-nineteenth century in a GIS-based study todetermine which broadscale environmental andsocioeconomic factors are most responsible for deter-mining where woodlands remained during the peakof agricultural land clearance. As with modern land-scapes, they found that woodlands were more likelyto be found farther from mid-nineteenth-centurybuildings and roads. Buildings and roads in turntended to be closer to the coast, near fresh water, andat lower elevations.

Historical descriptions of seventeeth-centurycoastal New England

To complement this effort, John Burk gathered seven-teenth-century historical data for the coastal region aspart of a broad effort, including paleoecologial andarchaeological studies, to understand vegetation com-position, structure, and dynamics of the area prior toEuropean settlement. Some 230 primary sources, cov-ering the region from Long Island to Cape Cod andthe North Shore, were researched, yielding 550 dis-tinct landscape and vegetation descriptions with vary-ing length and detail. Explorer accounts checked inthe past year included the voyages of Henry Hudson, Thomas Dermer, and Samuel de Champlain.The papers of Colonial figures such as RogerWilliams, John Winthrop, Thomas Morton, John Jos-

REU student Alex Ireland and Wyatt Oswald.

Tony D’Amato next to a 489-year-old eastern hemlock inan old-growth forest within Mohawk Trail State Forest

25

selyn, Alexander Hamilton, and others provided infor-mation about native populations and lifestyles, trade,and travel descriptions, as did Indian war journals ofJohn Mason, John Underhill, and Benjamin Church.

A search of town, state, and colony records wasconducted as a follow up to the explorer accounts,with initial emphasis on geographic areas notaddressed in the former, i.e., areas of Connecticut andRhode Island. The town records included ordinancesto burn woods, uplands, and underbrush; citations ofwoodlands to be divided by proprietors, depletion ofwood by liming and mill operations; and locations ofgrasslands, meadows, Indian fields, and sawmills.Every town checked enacted some sort of timber con-servation act by the early 1700s, with many citingconsiderable losses of forest. A number of towns,including those on Cape Cod, also passed acts to pro-tect and/or grant permission to “box,” “milk,” or “tur-pentine” pine trees. Similar citations, on a wider geo-graphic scale, were found in the Massachusetts Bay,Plymouth Colony, Rhode Island, Connecticut, andNew York Colonial government records.

Historical ecology of southwestern Connecticut

As part of the Highstead Arboretum project, JohnBurk and Brian Hall researched historical recordsrelated to the town of Redding and Fairfield County,Connecticut at a variety of repositories. To date, adataset of over 500 witness trees has been compiledfrom deed surveys in the town land records to 1800,and other relevant sources such as historical maps,

diaries, farm journals, town records, photographs,and publications have also been checked and enteredinto a comprehensive bibliography. These trees willbe summarized for each of the ten districts in Red-ding, allowing us to describe spatial variation inearly-settlement forest tree species composition.

Brian Hall has also gathered a wide variety ofdigital datalayers and historical maps to utilize in aGIS. Historical atlases will be used to documentchanging patterns in roads and houses since 1857,and determine the locations of wood-intensive indus-tries. Maps of surficial and bedrock geology will helpcharacterize the town’s environment while GIS layersfrom the University of Connecticut will help us iden-tify patterns of landcover change since the 1980s.

HARVARD FOREST ECOLOGY SYMPOSIUM 2005 (*DENOTES SUMMER STUDENTS)

M. Albani, A. M. Ellison, D. M. Medvigy, P. Moorcroft, and D. A. Orwig. Modeling HWA Impact at the Region-al Scale with the Ecosystem Demography Model.

J. E. Anderson, J. Blair, R. Dubayah, M. Hofton, P. Hyde, R. Knox, M. Martin, B. Peterson, and M. Smith.The Use of Waveform Lidar to Measure Temperate Mixed Deciduous Forest.

M. Bank and R. I. McDonald. Landscape-level Effects of Forest Harvesting on Vascular Plant Composition in Massachu-setts.

A. Barker Plotkin, A. M. Ellison, D. R. Foster, and D. R. Orwig. Logging in the Hemlock Manipulation Study.A. Barker Plotkin and D. R. Foster. The Tom Swamp Mapped Overstory Plots, 1990–2003.E. Boose, P. Barten, and B. Colburn. Prospect Hill Hydrological Stations.J. Burk, D. R. Foster, and G. Motzkin. Early Historical Records of the Northeast Coast.P. E. Busby, D. R. Foster, and G. Motzkin. Stand Dynamics and Vegetation History of Beech Forests on Naushon Island,

MA.J. Butler and A. M. Ellison. Nitrogen Uptake and Translocation in Sarracenia purpurea.J. Butler, A. M. Ellison, and N. J. Gotelli. Cycling of Inorganic Nitrogen through the Pitcher Plant Food Web.H. S. Callahan, J. Aber, K. del Fierro, A. E. Patterson, and H. Zafar. Impacts of Chronic N Amendments on Tree

Reproduction in an Oak-Dominated Stand at Harvard Forest.B. Choat, M. Holbrook, and E. C. Lahr. The Spatial Pattern of Air Seeding Thresholds in Mature Sugar Maple Trees.B. Colburn, M. Bank, E. Boose, D. R. Foster, and D. A. Orwig. Predicting How Changes in Forest Stand Composi-

tion Will Alter Stream Ecosystems at Multiple Spatial and Ecological Scales.B. M. Collins*, B. Colburn, and B. Sobczak. Subsurface Flowpaths Drive Spatial Variation in the Macroinvertebrate

Community of an Intermittent Headwater Stream in Central New England.T. D’Amato and D. A. Orwig. The Structure, Composition, and Dynamics Of Old-Growth Forests in the Berkshire Hills

and Taconic Mountains, Western Massachusetts.B. Dail, J. Aber, E. Davidson, and D. Hollinger. Seeing the Forest for the Soils: Fate of a 15N Foliar N Addition in a

Mature Spruce-Hemlock Stand, Howland, Maine.M. J. Daley and N. Phillips. Interspecific Variation in Nighttime Transpiration and Stomatal Conductance.E. Davidson, K. Savage, and S. Trumbore. Decadal-Scale Measurements of Decadal-Cycling Forest Soil Carbon.E. Davidson, B. Dail, D. Hollinger, A. Richardson, and K. Savage. A Distinct Seasonal Pattern of the Ratio of Soil

Respiration to Total Ecosystem Respiration in a Spruce-Dominated Forest.B. G. DeGasperis, D. R. Foster, and G. Motzkin. Effects of Historical and Modern Anthropogenic Disturbances on the

Distribution and Demography of Berberis thunbergii (Japanese barberry).J. W. Elkins, G. S. Dutton, E. Gottlieb, M. B. McElroy, and S. Wofsy. A Harvard Forest Perspective: Are EPA U.S.

Emissions of the Greenhouse Gases Correct.E. Faison and D. R. Foster. The Effects of Moose and Deer on Harvested Oak-Pine Forests of Central Massachusetts.J. Hadley and P. Kuzeja. Water Use by Eastern Hemlock (Tsuga canadensis) Forest and Deciduous Forests in Central Mass-

achusetts: Hydrologic Implication. R. Hanifin*, J. Melillo, and J. Mohan. First-Year Reproductive Responses of Two Herbaceous Species to Experimental Soil

Warming at Harvard Forest.S. Jefts and D. A. Orwig. The Effects of HWA Outbreaks on Ecosystem Level Changes in Southern New England.J. Jenkins, K. McKnight*, and G. Motzkin. The Flora of Harvard Forest.C. Jones, D. Gonzalez-Kreisberg*, L. Hutyra, K. McKain*, W. Munger, E. Pyle, and S. Wofsy. Carbon Exchange

Studies at Harvard Forest Using Eddy Flux Tower Measurements and Ground-based Ecological Measurements.J. Karagatzides, J. Butler, and A. M. Ellison. Sarracenia Can Directly Acquire Organic Nitrogen and Short-Circuit the

Inorganic Nitrogen Cycle.D. Kittredge and K. Grogan*. The Parcelization of Forests and Timber Harvest.C. Lai and J. Ehleringer. Carbon Isotope Ratio of Nighttime Respiration, Leaf, and Soil Organic Matter in a Temperate

Deciduous Forest.W. H. Liu, A. V. Bright, J. Budney, D. Curran, B. C. Daube, L. Hutyra, C. Jones, W. Munger, E. Pyle, S. R.

26

Saleska, and S. Wofsy. The Effects of Selective Management on Carbon in Woody Debris: Measurements of Respira-tion and Environmental Conditions.

A. Magill and J. Aber. Chronic Nitrogen Amendment Study.B. Mathewson, B. Colburn, and D. R. Foster. Eastern Redback Salamander (Plethodon cinereus) and Juvenile Eastern

Red-Spotted Newt (Notophthalmus viridescens) Abundance in Eastern Hemlock-Dominated Stands and Mixed DeciduousStands.

K. A. McKinney, H. A. Fuchs, and A. Lone. Fast Time Response Measurements of Volatile Organic Compound (VOC)Concentrations and Fluxes at Harvard Forest.

Q. Min. A Regional Climatology of Cloud and Aerosol for Forest-Atmosphere Exchange.R. Minocha and S. L. Long. Effects of Nitrogen Fertilization/Deposition on the Foliar Nutrition and Stress Physiology of

Coniferous and Broadleaf Trees Across Northeastern US.J. Mohan, J. H. Blanchard, E. Burrows, J. Melillo, and P. Steudler. Soil Warming at Harvard Forest: Effects on Bio-

geochemistry and Forest Trees.P. Moorcroft, M. Albani, D. M. Medvigy, and S. Wofsy. Constraining the Exchange of Carbon, Water, and Energy

Between North-Eastern Ecosystems and the Atmosphere: Results From a Regional-Scale Coupled Terrestrial BiosphereModel.

J. O’Keefe. Woody Species Phenology, Prospect Hill Tract, Harvard Forest — 2004.J. O’Keefe and P. Snow. Regeneration Following Clearcutting of Red Pine verstory — Year 15.D. A. Orwig, L. Barbash, D. R. Foster, and H. Lux. Community and Ecosystem effects of HWA-induced Logging.D. A. Orwig, P. Del Tredici, D. Foster, H. Lux, and R. Schulhof. Comparison of Hemlock Cutting Following HWA

Infestations in Urban vs. Rural Forests.D. A. Orwig, M. Manner*, D. Niebyl*, and N. Povak*. Landscape Level Analyses of Hemlock Woolly Adelgid Out-

breaks in Massachusetts.W. Oswald and D. R. Foster. Lake-Sediment Evidence for Late Holocene Climatic Variability Across Southern New Eng-

land.W. Oswald, D. R. Foster, and G. Motzkin. Long-Term History of Vegetation and Fire in the Pine Barrens Region of Long

Island, New York.W. Oswald, E. Faison, and D. R. Foster. Vegetation Dynamics Across Southern New England During the Middle

Holocene.N. Pederson, A. Barker Plotkin, G. C. Jacoby, E. Pyle, and S. Wofsy. A Regional Context for Northern Red Oak

Growth Rates at the Harvard Forest.N. Phillips and M. J. Daley. Dynamic Coupling of Stomata and Tree Hydraulics.F. E. Rockwell and D. Hollinger. The Contribution of Gas-Filled Fibers to Positive Sap Pressures in Sugar Maple.N. A. Scott, B. Dail, E. Davidson, D. Hollinger, H. Hughes, J. T. Lee, and C. Rodrigues. Recovery of Net Ecosys-

tem Carbon Sequestration Following a Shelterwood Harvest at Howland Forest, Maine, U.S.A.S. Sevanto, M. Holbrook, W. Munger, and S. Wofsy. Diurnal Stem Diameter Variations: A New Tool for Detecting

Sap Flow.B. Stadler, T. Muller, and D. A. Orwig. The Ecology of Energy and Nutrient Fluxes in Hemlock Forests Invaded by Hem-

lock Woolly Adelgid. K. Stinson, L. Durbin*, S. Kaufman, and F. Lowenstein. Responses of a New England Forest Community to Increas-

ing Levels of Invasion.K. Stinson and J. Klironomos. Exotic Plant Invasion Degrades Local Mycorrhizal Association, Alters Community Succes-

sion and Limits Restoration.K. Stinson, C. Chang*, and K. Donohue. Effects of Varying Environmental and Maternal Habitats on the Performance,

Demographic Structure, and Population Dynamics of Alliaria petiolata.E. T. Sundquist. Monitoring and Modeling of Soil Heat, Water, and Gas Transport.R. K. Varner and P. M. Crill. Two Years of High Temporal Frequency Measurements of CO2 Efflux from Soil. B. Von Holle. Ecosystem Effects and Legacies of the Introduced N-Fixing Tree, Robinia Pseudoacacia, in the Upland Coastal

Forests of Cape Cod, MA.H. Wu and X. Lee. Response of Soil Respiration to Rain in a Temperate, Hardwood Forest in Massachusetts.

27

HARVARD FOREST ECOLOGY SYMPOSIUM 2006 (*DENOTES SUMMER STUDENTS)

A. Barker Plotkin, D. R. Foster, and N. Levy*. Coarse woody debris dynamics after hemlock removal.J. Blanchard, F. Bowles, E. Burrows, H. Lux, J. Melillo, J. Mohan, and P. Steudler. Barre Woods Soil Warming

Project: Effects on biogeochemistry and forest trees after three years of warming.E. Boose, J. Hadley, and S. Singh*. Preliminary water budget for Prospect Hill Tract.T. Brodribb and N. M. Holbrook. Leaf hydraulic conductance is highly variable and sensitive to leaf water potential.J. Burk, D. R. Foster, and G. Motzkin. “The Paradise of these Parts:” 17th century primary accounts of the northeast

coastal landscape.P.. Busby, C. Canham, D. R. Foster, and G. Motzkin. Tree response to hurricane disturbance in coastal New England.J. Butler and A. M. Ellison. Nitrogen cycling dynamics in the northern pitcher plant, Sarracenia purpurea.B. Choat, J. Anderson, N. M. Holbrook, and M. Zwieniecki. Dynamic changes in xylem hydraulic resistance via mod-

ulation of xylem sap ionic composition.R. Cobb and D. A. Orwig. Vegetation driven decomposition changes resulting from hemlock woolly adelgid.A. D’Amato and D. A. Orwig. The structure, composition, and dynamics of old-growth forests in the Berkshire Hills and

Taconic Mountains, western Massachusetts.M. Daley, J. Hadley, J. Pettijohn, and N. Phillips. A comparison of transpiration in black birch and eastern hemlock

stands.E. Davidson and K. Savage. Reconciling landscape-scale measurements of soil respiration with estimates of total ecosystem

respiration.B. DeGasperis and G. Motzkin. Being in the right place at the right time: a historical perspective on Japanese barberry

invasion.D. Diaz* and N. Tuross. Nitrogen isotope evidence for ecological changes and adaptations relating to agricultural land-use

legacies in the foliage of four New England plant species.A. M. Ellison. The Hemlock Removal Experiment: Initial changes in environmental variables following treatments.A. M. Ellison, E. Boose, L. Clarke, D. R. Foster, J. Hadley, L. Osterweil, and A. Wise. Ensuring reliable datasets

for valid environmental models and useful environmental forecasts.E. Faison, D. R. Foster, and G. Motzkin. Moose foraging ecology at their southern range boundary: the temperate forests

of central Massachusetts.E. Farnsworth and A. M. Ellison. Prey availability directly affects physiology and nutrient allocation in ten carnivorous plant

species.D. Fitzjarrald and A. Tsoyref. Forest-atmosphere exchange processes related to regional carbon budgets.S. Frey, A. Contosta, and M. Knorr. Microbial responses to soil warming and nitrogen additions.J. Jenkins, K. McKnight*, and G. Motzkin. The flora of Harvard Forest.C-T. Lai and J. Ehleringer. Interannual variability of carbon isotope ratios of ecosystem respiration in Harvard Forest between

2001-2005.H. Lux, P. Del Tredici and D. A. Orwig. Ecosystem responses to forest cutting after HWA infestation: New insights from

an urban site.B. Mathewson, B. Colburn, and D. R. Foster. Differences in eastern red-backed salamander populations in eastern hem-

lock forests and mixed deciduous forests.R. McDonald, D. R. Foster, and G. Motzkin. Land-use legacies, present-day forest fragmentation and harvesting, and

invasive species.K. McKain*, J. Bubier, E. Hammond Pyle, and S. Wofsy. Carbon accumulation at the Harvard Forest: A comparison

of methods for measuring tree biomass for regional extrapolation of the eddy-flux tower footprint.D. Medvigy, P. Moorcroft, and S. Wofsy. The state of the regional carbon cycle: Results from a constrained coupled ecosys-

tem-atmosphere model.Q. Min, D. Fitzjarrald, and S. Wang. Clouds, a controlling factor in terrestrial carbon uptake.J. Munger, C. Barford, J. Budney, D. Curran, E. Hammond Pyle, and K. McKain. Accelerating carbon uptake

rates at the Harvard Forest environmental measurement site.J. O’Keefe. Woody species phenology, Prospect Hill Tract, Harvard Forest – 2005.

28

S. Ollinger, J. Jenkins, M. Martin, J. Munger, L. Plourde, and M-L. Smith. Canopy nitrogen and forest carbonassimilation: Validation of local patterns and broad-scale satellite estimates.

D. A. Orwig, D. Niebyl*, and N. Povak*. Landscape level analyses of hemlock woolly adelgid outbreaks in Massachu-setts.

D. A. Orwig, L. Barbash, D. R. Foster, and H. Lux. Community and ecosystem effects of HWA-induced logging.W. Oswald and D. R. Foster. Late-Holocene history of coastal ecosystems of New England and New York.A. Paradis and J. Elkinton. Population dynamics of the hemlock woolly adelgid.J. Pettijohn and M. Daley, N. Phillips, and G. Salvucci. Evaluating the sensitivity of Acer rubrum L. sap flux to envi-

ronmental stress under two root-zone soil water limitation scenarios.E. Preisser, J. Elkinton, A. Lodge*, and D. A. Orwig. Range expansion and community impact of the invasive herbi-

vores Adelges tsugae and Fiorinia externa in New England.S. Record and A. M. Ellison. Seed Dispersal: Implications for forest regeneration. S. Sevanto, M. Daley, N. M. Holbrook, J. Pettijohn, and N. Phillips. Estimating sap flow using diurnal stem diam-

eter variations.T. Sipe and S. Strouse. Herbaceous stratum effects on nearground enriched CO2 profiles.B. Sobczak, E. Boose, B. Colburn, and J. Hadley. Examination of terrestrial and aquatic ecosystem linkages at Harvard

Forest.P. Steudler. Carbon monoxide uptake kinetics in unamended and long-term nitrogen amended temperate forest soils.K. Stinson, J. Klironomos, and B. Wolfe. The enemy of my friend: an invasive plant disrupts native tree seedling —

mycorrhizal fungi mutualism.M. Trumbull*, A. Barker Plotkin, and D. R. Foster. Understory response over 15 years following simulated hurricane

at the Harvard Forest.R. Varner and P. Crill. Three years of high frequency measurements of soil surface CO2 efflux at Harvard Forest.J. Weishampel, J. B. Blair, A. Cooper, J. Drake, D. R. Foster, M. Hofton, and G. Motzkin. Forest canopy recov-

ery from the 1938 hurricane and subsequent salvage damage measured with airborne LiDAR.B. Wolfe, J. Klironomos, A. Pringle, V. Rodgers, and K. Stinson. Impacts of the invasion of garlic mustard (Alliaria

petiolata) on forest fungi.

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30

BULLARD FELLOWS

John Briggs (Arizona State University and the Cen-tral Arizona Phoenix LTER site) collaborated withDavid Foster, Aaron Ellison, and Emery Boose, toexplore use of historical data to help explain andunderstand modern day ecological patterns and inparticular plant distributions. John applied theseskills to help analyze data that he has collected forover twenty years at the Konza Prairie Biological Sta-tion (a LTER research site since 1981). John alsoworked on manuscripts and on finishing a book TheBiology of Grasslands (Oxford University Press) that hehad recently started with Alan Knapp (ColoradoState University) and Scott Collins (University ofNew Mexico).

Conghe Song’s (University of North Carolina)research was primarily focused on remote sensing ofvegetation and ecological modeling. He investigatedthe potential of extracting forest canopy structurefrom high-resolution optical remotely sensed imagerybased on field data collected during the summer of2005. On ecological modeling, he has linked hisMVP model with Farquhar’s photosynthesis modeland Leuning’s stomatal conductance model to understand the importance of canopy structure onradiation interception, latent heat transfer, and carbon assimilation of forest ecosystem. In addition,he collaborated with a colleague in China to studythe forest cover change in China from 1949 to 2003 as a result of afforestation, deforestation, andreforestation.

Richard D. Bowden (Allegheny College) spenthis sabbatical year working on projects related to

long-term and current anthropogenic alterations toforest C and N dynamics. Continuing a Collabora-tive Research at Undergraduate Institutions projectbegun in 2000 with Tim Sipe (Franklin and MarshallCollege) and Chuck McClaugherty (Mt. Union Col-lege), Bowden continued investigations into the long-term influences of historic agriculture on current for-est ecosystem patterns and processes. Building on thestrong history of land-use research at the forest, Richand his colleagues investigated vegetation and soil inforests that had been formerly cultivated, pastured,or left as woodlots, and which were then releasedfrom agriculture during the height of agricultural pro-duction in the mid-1800s.

Richard D. Bowden.

In this study, despite nearly two centuries of for-est regrowth, soils show profound differences amongthe former land-use sites. For example, total soil car-bon is much lower in the former agricultural sitesthan in the former woodlots. This information is crit-ical because up to 85 percent of the regional land-scape has been either plowed or pastured during thenineteenth-century agricultural era, and becauseunderstanding soil stores of carbon is important inquantifying links between the global carbon cycleand global climate change.

Rich also continued work on the Detritus Inputand Removal Treatment (DIRT) project. Collaborat-ing with colleagues at Debrecen University in Hun-gary, he examined soils at the Sikfokut (Hungary)International LTER DIRT site.

Since beginning her Bullard Fellowship in Febru-ary 2005, Elizabeth Farnsworth (New England WildFlower Society) has completed two manuscripts ana-lyzing a large data set on dozens of rare plant speciesin New England. She finished writing and illustratingher book (with Cheryl Lowe), the Peterson Field Guideto Ferns and Their Related Families. With co-authorsJohn and Wendy Sinton, Elizabeth began research ona guide to the Connecticut River.

She also collaborated with Aaron Ellison andREU student Cheryl Hester, from Pueblo Commu-nity College, on an experiment examining nutrientlimitation and photosynthetic physiology of tenspecies of carnivorous plants. Elizabeth gave research

talks at Harvard Forest, Wesleyan University, East-ern Connecticut State University, and Bates Collegeand has led/co-led two workshops in scientific ethicsfor Harvard Forest staff and REU students.

Julia Jones (Oregon State University) completeddrafts of five articles, became an editor for Ecology,and completed a draft of a book intended as a grad-uate text, entitled Spatiotemporal Statistics in Ecology andEarth Science. In collaboration with Aaron Ellison, sheapplied spatiotemporal analysis techniques to lookfor patterns in tree species in pollen cores that DavidFoster and his group have amassed. Julia’s analysis oflong-term streamflow records from Harvard Forest’s“sister” LTER sites, such as Hubbard Brook, Cowee-ta, and the Andrews has proven relevant to the plansfor establishing stream gaging at Harvard Forest.

Greg Jordan (University of Tasmania, Hobart)investigated the plant macrofossil flora from StonyCreek Basin, a remarkable fossil assemblage providesa continuous record of vegetation and insect faunasspanning some 300,000 years (approximately 1.8–1.5million years ago), as dated using radiometric andsedimentological methods.

Greg’s work involved scanning electronmicroscopy and comparative analyses to identify sev-eral thousand specimens from approximately eightyspecies of plants. The sampling covered depth rangeof approximately four meters from the upper part ofthe section, and corresponded to a full climatic cycle.

David Lee, (Florida International University)completed a book manuscript and illustrations,which are now in review by University of ChicagoPress. With John O’Keefe, David completed a surveyof leaf senescence color and anthocyanin productionin herbaceous plants at the forest (sixty species). Hecreated and installed autumn foliage website(http://harvardforest.fas.harvard.edu/research/leaves/autumn_leaves.html) with Julie Pallant and JohnO’Keefe, and conducted pilot measurements on thespectral distribution of radiation within leaf traps andreflected from outside, in Sarracenia leucophylla and S.alata.

Gidon Ne’eman (Department of Biology, Uni-versity of Haifa-Oranim, Israel), assisted by his wifeRina (Anne Frank High School, Branco Weiss Insti-tute, Kibbutz Sasa, Israel) collaborated with AaronEllison to study pollinator-prey conflict in the carniv-orous pitcher plant Sarracenia purpurea.

The pitchers and flowers of S. purpurea both aremostly red and produce nectar, traits which attractpollinators to flowers and prey to pitchers. Therefore,

31

Elizabeth Farnsworth.

it has been suggested that pitchers may mimic flow-ers to capture prey, potentially resulting in a prey-pollinator conflict; however this conflict has neverbeen experimentally tested.

The results indicated that none of the major pol-linators of S. purpurea was found captured within thepitchers, indicating that the flowers and pitchersattract different organisms, minimizing competitionas well as the possible existence of a prey-pollinatorconflict.

Rina Ne’eman, in collaboration with the Wyatt

Oswald and Elaine Doughty, prepared a collection ofpollen of all Sarracenia species growing in HarvardForest greenhouse. With Aaron Ellison she also per-formed phenological observations and measurementsto determine the degree of temporal and spatial sep-

aration between the flowers, production of new pitch-ers, and senescence of old pitchers in all Sarraceniaspecies growing in the Harvard Forest greenhouse.

Peter Thomas (Keele University, UK) completeda textbook on forest ecology, a joint project with John Packham of the University of Wolverhampton,UK, to be published by Cambridge University Press.As a result of his time in Petersham, the book has astrong Harvard Forest flavor. He also finished apaper on a sixteen-year experiment on grassland fireson cacti in Arizona (published in Plant Ecology), andwrote two popular science articles on forests for theJournal of the International Tree Foundation and TheDendrologist. In between he rewrote a chapter of aprevious book Trees: Their Natural History with an

eye to a second edition, and used the excellentlibraries to work on a monograph on juniper for theBiological Flora of the British Isles series in Journal ofEcology, cowritten with colleagues at Keele and Gary-ounis University, Libya. His wife, Judy, thoroughlyenjoyed voluntary work at the Petersham elementaryschool, and their younger son experienced the cul-ture shock of attending Mahar Regional High Schoolfor a semester.

Jill Thompson (Luquillo LTER site, Universityof Puerto Rico) enjoyed a wonderful year and all theseasons at Harvard Forest. During the year she con-tinued her research on the effects of land-use historyand hurricane damage on the subtropical wet forestin Puerto Rico using census data from the LuquilloForest Dynamics Plot (LFDP). She also worked on avariety of papers about the LFDP concerning alienspecies dynamics, orchid distribution, methods for

32

Gidon Ne’eman.

Rina Ne’eman.

Peter and Judy Thomas.

measuring biodiversity, nonrandom processes anddiversity in tropical forests, effects of hurricane dam-age on light availability, seedling recruitment in hur-ricane driven forest, Bayesian analysis of hurricanedamage and tree survival, phylogenetic patterns intropical forest, and tropical wet forest versus riparianvegetation. Jill gave a seminar on her research in trop-ical forest, a workshop about using relational data-bases for data management, explained canopy fish-eye photography analysis to Kristina Stinson andAnna Bellafiore, spent a week helping Posy Busbycore trees on Naushon Island, and taught the sum-mer REU students Scottish country dancing.

EDUCATIONAL ACTIVITIES

Betsy Colburn mentored MFS student Brooks Math-ewson in his research on terrestrial salamanders inhemlock and deciduous forest stands. She alsoreviewed the master’s thesis of Holly Jensen (AntiochNew England Graduate School), in which Holly pre-sented the results of surveys carried out on headwa-ter streams at Mt. Wachusett; advised (informally)graduate students from Wheaton College (IL),UMass, and Clark University; and judged studentpapers at the annual meeting of the North AmericanBenthological Society.

Each spring Glenn Motzkin, John O’Keefe, andDave Orwig joined David Foster in leading the Har-vard Forest Freshman Seminar. The Seminar meetsover four weekends and provides an overview ofHarvard Forest research and an introduction to the

landscape and history of New England. It concludeswith each student undertaking an independent study.

P. Barry Tomlinson taught two courses at theKampong Garden of the National Tropical BotanicalGarden in Coconut Grove, Miami, Florida. TheKenan College Professors Course June 12–24demonstrated teaching methods in tropical botany sothat instructors could increase the coverage of botanyin introductory biology courses. “Biodiversity ofTropical Plants,” was held June 26–July 21. It is aHarvard Summer School course for graduate stu-dents, which expanded on existing botanical knowl-edge in a tropical environment.

Serita Frey (University of New Hampshire) pre-pared a dataset for publication in Teaching Issues andExperiments in Ecology (http://tiee.ecoed.net/) usingdata collected over the past fifteen years at theChronic Nitrogen Addition Study. The activityguides students to ask and address questions aboutthe long-term effects of chronic N additions on netprimary productivity, tree photosynthetic capacity,soil respiration, and soil microbial biomass in two for-est types. Students examine the data, ask their ownquestions, and then propose and prepare the mostmeaningful figures to get at these questions.

Summer Research Program

The Harvard Forest Summer Student Research pro-gram, coordinated by Edythe Ellin and assisted byHilary Crowell attracted a diverse group of 22 stu-dents to receive training in scientific investigations,and to gain experience in long-term ecologicalresearch. All students work closely with researcherswhile many conduct their own independent studies.The program includes weekly seminars from residentand visiting scientists, discussions on career issues inscience, and field exercises on soils, land-use history,and plant identification. An annual field trip is madeto the Institute of Ecosystem Studies in Millbrook,NY, to participate in a Forum on Careers in Ecology.Students present major results of their work at theAnnual Summer Student Research Symposium inmid-August.

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Jill Thompson.

Summer Students 2005

Emily Austin Hampshire CollegeRyan Barba Assumption CollegeCharles Boyd Centre CollegeDaniel Brese Bennington CollegeLaura Briscoe College of the AtlanticJonah Butler New College of FloridaSusan Cheng Columbia UniversityRobin Collins University of FloridaAntoinnine Cooper Oakwood CollegeDavid Diaz Harvard CollegeCheryl Hester Pueblo Community CollegeKristin Ivy Grambling State University Daniel Katz Bard CollegeMatthew Kaufman Keene State CollegeNicholas Kuzma Franklin & MarshallPhil Labranche Holyoke Community

CollegeKatherine Lenoir Wellesley CollegeBennet Leon Bates CollegeNatalie Levy University of

California–Berkeley

Alexandra Lodge Kenyon CollegeKathleen Logothetis Franklin & MarshallJennifer McGinnis Cornell UniversitySafina Singh Mount Holyoke CollegeJens Stevens Carleton CollegeStephanie Strouse Franklin & MarshallSarah Truebe Stanford UniversityMathew Trumbull Hampshire CollegeLinh Vuong University of Puget SoundKelly Walton State University of

New YorkKirsten Ward Brigham Young UniversityBrian Warshay Cornell UniversityMelissa Whitaker Prescott CollegeMarit Wilkerson University of Texas, AustinJames Willacker, Jr. State University of

New YorkGrace Wu Pomona College

Assistant Program Coordinators

Kathryn McKainThomas Mulcahy

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Summer Students 2006

Alex Arguello St. Mary’s College of Maryland

Laura Briscoe College of the AtlanticKevin Burls Mount Union CollegeStephanie Day Howard UniversityJustin Frisino SUNY Albany

University of AlbanyDana Graef Princeton UniversityAlison Grantham Mount HolyokeAlex Ireland Clarion UniversityFrances O’Donnell Harvard UniversityRebecca Orozco University of KansasRachel Osborn Hampshire CollegeChristine Rollinson Oberlin CollegeTimothy Rowell College of the Holy Cross

Lisa Schauer New Mexico StateUniversity

Kaya Schmandt Brown UniversityStephanie Searle Columbia UniversityBrynne Simmons Johnson C. Smith

UniversityLori Simpers Susquehanna UniversitySafina Singh Mount Holyoke CollegeRachel Stahr Hampshire CollegeChelsea Vario University of New

HampshireJessica Megan Woltz North Carolina State

University

Assistant Program Coordinator

Hilary Crowell

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Master of Forest Science Program

In Spring 2006, four students graduated from theHarvard Forest MFS program. Posy Busby, thesistitle: Beech dynamics and dominance in coastal New Englandforests. Brian DeGasperis, thesis title: Windows of oppor-tunity: historical and ecological insights into plant invasions.Edward Faison, thesis title: Moose foraging in the temper-ate forests of central Massachusetts: a natural re-wildingexperiment. Brooks Mathewson, thesis title: A compari-son of eastern red-backed salamander populations in hemlock-dominated and mixed deciduous forests in north central Mass-achusetts.

Building the LTER Schoolyard Program

With funding from NSF’s Schoolyard (sLTER) andEd En Venture Fund programs, along with the Mass-achusetts Environmental Trust, we were able to sup-port twenty-seven teachers in implementing fieldresearch projects in their schoolyards. Pam Snow,environmental educator, led this project with JohnO’Keefe. In this effort, we continued our collabora-

tions with the Hitchcock Center for the Environmentin Amherst and the Millers River EnvironmentalCenter in Athol, while formalizing our partnershipwith the Nashua River Watershed Association. Wewere also able to bring this schoolyard model west toschools in the Deerfield River Watershed. To reachthese schools, we collaborated with the Student Con-servation Association Mass. Parks AmeriCorps,based in Hawley, Massachusetts.

Harvard Forest ecologists Betsy Colburn, DavidOrwig, and John O’Keefe continued to help trainteachers in implementing freshwater and forest ecol-ogy protocols modified for school use. This training

included a two-day Summer Institute and two day-long workshops at Harvard Forest during the schoolyear. Ecologists, environmental educators, and ourinformation manager, Emery Boose, assisted in theseformal trainings at the Harvard Forest. Water-levelfluctuations in streams and vernal pools were one ofthe subjects studied by teachers. Betsy Colburnworked with Emery Boose and Pam Snow on meta-data and data formatting, and with teachers and Pam

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Posy Busby, Brooks Mathewson, Ed Faison, and Brian DeGasperis.

on data analysis. Each participant was also coachedby an environmental educator on site at their schoolas needed throughout the school year. Dr. Boose wasable to make a host of data-management documentsand tools available to teachers on our HF Web sitethis year. Among the Web postings are two years ofstudent data from participating schools. We expect tocontinue to post student data for future year’s proj-ects and eventually develop a long-term dataset ofstudent data.

As a result of these professional developmentefforts, not only did teachers successfully implementfield ecology protocols, but three teachers also wenton to train fellow teachers, including these protocolsin their presentations. Kate Bennett and MaryGagnon of Briggs Elementary School in Ashburn-ham recommended use of HF protocols to teachersattending the Massachusetts Environmental Educa-tion Society’s annual conference. Michael Silverstoneof Wildwood Elementary School in Amherst sharedthe protocols as a teaching tool for teachers attendinga workshop in strategies for English Language Learners.

Elaine Senechal of Tewksbury High Schoolspread the news of our program to the general pub-lic by contacting a Boston Globe reporter, who pub-lished a full-length story about their schoolyard proj-ect along with color photos and a cartoon. Our pro-gram was also featured in a number of community

newspapers in our region, featuring teachers andcoaches from Athol, Ashburnham, Warwick, andTewskbury.

In an effort to strengthen our connections to thenational LTER education community, Pam attendeda national LTER Education/Children’s Book Com-mittee meeting in September 2005 and presentedHarvard Forest’s “Buds, Leaves, and Global Warm-ing” schoolyard protocol at a teacher institute led byPIE-sLTER representative, Liz Duff.

Harvard Forest has once again been awardedfunding from the NSF sLTER program to allow usto continue to support some teachers in implement-ing field ecology studies at their schools during FY07.However, the Ed En Venture program, which pro-vided the bulk of our funding in the past two years,was not offered this year. Fortunately, we havereceived a three-year grant from the Cardinal BrookTrust to help support our local schoolyard projects.We will continue to work with the Millers RiverEnvironmental Center and the Nashua River Water-shed Association to support them in their efforts toinvolve teachers in Warwick, Athol, and Ashburn-ham in our schoolyard field studies, with the supportof the Massachusetts Environmental Trust.

ACTIVITIES OF THE FISHER MUSEUM

The Fisher Museum plays an important role in theeducational mission of the Harvard Forest by pro-viding a public outlet for research in forest biology,conservation, and management. The Museum alsoprovides a unique setting for conferences and work-shops sponsored by the Forest and outside organiza-tions. Dr. John O’Keefe has primary responsibilityfor the development of activities and coordination ofthe use of the Museum and he is assisted by PamSnow.

Our great group of Museum volunteers continueto allow the Fisher Museum to provide a very muchappreciated weekend schedule from May throughOctober. Each November the group shares food, fel-lowship, and ideas at our traditional VolunteerRecognition Dinner. In 2005, Bob Clark, WaltDavidson, and Bob Lane received special recognitionfor being the most active volunteers during the sea-son. In 2006 Bob Clark was again recognized forbeing the most active volunteer during the season. At each dinner, volunteer coordinator Mary AnnWalker received sincere thanks for her tireless andenthusiastic work to keep the program running

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Budding ecologists.

smoothly. The group has been deeply saddened bythe death of two of its founding members, PeterStrong during the winter of 2004–05, and HectorCameron the following winter. A memorial servicefor Peter, Mary Ann’s husband, was held in theMuseum on Saturday, February 5, 2005. Peter’seclectic knowledge, recitations, and twinkle will besincerely missed.

During 2004–05 the Museum provided pro-grams for twenty-five elementary and secondaryschools, thirty-five college and university classes, andthirty-eight community and professional groups.During the summer the Museum hosted two groupsof inner-city youth in our continuing collaborationwith the University of Massachusetts Extension“Learn About Forests” program. Each group spentthe day exploring, learning, and working on theSchwartz lot and then enjoyed a cookout behindShaler Hall.

During 2005-06 the Museum provided programsfor thirty-two elementary and secondary schools,fifty-three college and university classes, and twenty-nine professional and community groups. Althoughthese programs varied considerably depending uponeach group’s age, background, and areas of interest,each group learned about the land-use history of theregion and current research and conservation activi-ties at Harvard Forest.

Meetings, Conferences, Seminars 2004–05

In August the Museum and Forest hosted a three-dayworkshop on soil microbial communities andprocesses organized by Serita Frey, one of our collab-orators at the University of New Hampshire. In Jan-uary, in collaboration with the Harvard MedicalSchool Center for Health and the Global Environ-ment, we hosted a three-day meeting of the Interna-tional Agriculture and Human Health Task Force. InFebruary we hosted a workshop cosponsored by theNational Science Foundation LTER program net-work office and organized by Aaron Ellison on theimpact of removal of foundational species by pest andpathogens on the structure and dynamics of forestedecosystems. This workshop brought together repre-sentatives from several LTER sites studying similarprocesses to better coordinate cross-site research proj-ects. And in May the National Science Foundationand Woods Hole Oceanographic Institute cospon-sored a week-long meeting at Harvard Forest focus-

ing on the population biology of albatrosses. The Six-teenth Annual Harvard Forest Long-Term EcologicalResearch Symposium and National Institute forGlobal Environmental Change meeting was held inthe Museum on February 23.

Meetings, Conferences, Seminars 2005–06

In August the Museum and Forest hosted a weeklongseminar course on research methods organized byLynn Adler, a recently hired faculty member at theUniversity of Massachusetts. Over the winter theForest hosted a series of retreats, one for each cohortof Harvard University Organismic and EvolutionaryBiology (OEB) graduate students. In January andagain in April Harvard Forest hosted planning meet-ings for the National Science Foundation’s NEONinitiative and in May we hosted a joint retreat for agroup of Harvard Medical School and FAS faculty.The Seventeenth Annual Harvard Forest Long-TermEcological Research Symposium and National Insti-tute for Global Environmental Change meeting washeld in the Museum on April 12.

Other groups meeting at Harvard Forest includ-ed the Greater Worcester Community FoundationStudent Grant Workshop, Irish Foresters SocietyNorth American tour, Massachusetts Audubon Soci-ety, Massachusetts Barn Preservation Task Force,Massachusetts Chapter of the American ChestnutFoundation, Massachusetts Department of Conser-vation and Recreation, Massachusetts ExecutiveOffice of Environmental Affairs/Land Trust Retreat,Massachusetts Extension Service Coverts Program,Massachusetts Forestry Association, MassachusettsForestry Forum, Massachusetts Forest Steward’sTraining, Massachusetts State Forestry Committee,Mount Grace Land Conservation Trust, MountWachusett Community College, New EnglandForestry Foundation, National Tree Farm Conven-tion tour, Northeast Soil and Water ConservationAssociation, North Quabbin Regional LandscapePartnership, The Nature Conservancy, Quabbin toCardigan Conservation Collaborative, The Trusteesof Reservations, U.S. Fish and Wildlife Service andU.S. Department of Agriculture Natural ResourcesConservation Service.

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Speakers in the Harvard Forest Seminar 2005–2006 series included:

Lynn Adler, University of Massachusetts, Ecology and evolutionary consequences of attraction and defense in plant-animal interactions.

Robert Bertin, Holy Cross College, The flora of Worcester County: patterns and changes. John Briggs, Arizona State University and Harvard Bullard Fellow, Ecosystems in transitions: tallgrass prairie and

Sonoran desert.Posy Busby, Harvard Forest, Beech dynamics and dominance in coastal New England forests.Liza Knapp, University of Massachusetts, Correlates and components of growth in native and invasive woody plants.Charles Davis, Organismic & Evolutionary Biology, Harvard University, Phylogenetic studies in Malpighiales:

dating the origin of tropical rain forests, host-to-parasite gene transfer, and the geography of gene swapping.Brian DeGasperis, Harvard Forest, Being in the right place at the right time: a historical perspective on Japanese

barberry invasion.Brian Donahue, Brandeis University and Harvard Bullard Fellow, Husbandry was once a sacred art:

environmental history and conservation.Edward Faison, Harvard Forest, Moose foraging in the temperate forests of Central Massachusetts: a natural re-wilding

experiment.Mark Harmon, Oregon State University and Harvard Bullard Fellow, Developing a multi-scale perspective of

forest carbon dynamics.Martha Hoopes, Mount Holyoke College, Invasions on the horizon: the spread and impact of non-native species.Jerry Jenkins, Glenn Motzkin, and Kirsten Ward, Harvard Forest, The flora of Harvard Forest.Charles Lord and Eric Strauss, Urban Ecology Institute, The Urban Ecology Collaborative: an experiment in

cross-city research for urban communities.Brooks Mathewson, Harvard Forest, Differences in eastern red-back salamander populations in eastern hemlock-

dominated and mixed deciduous forests in north central Massachusetts.Mark McDonnell, University of Melbourne, Australia

The challenges and opportunities of studying the ecology of human settlements Down Under.Laura Meyerson, University of Rhode Island, Tracking nonnative species at the national and state level.Jacqueline Mohan, The Ecosystems Center, Marine Biological Laboratory

Global change impacts on eastern forests.Anne Pringle, Organismic & Evolutionary Biology, Harvard University, Last chance to know? Using clues from

the literature and a genome to understand the biogeography of the death cap mushroom Amanita phalloides.Richard Primack and Abe Miller-Rushing, Boston University, The impact of climate change on the birds and plants

of Massachusetts.Ferran Roda, Autonomous University of Barcelona, Land-use patterns, forest fragmentation and biodiversity in the

Barcelona metropolitan area. Dana Royer, Dept. of Earth & Environmental Sciences, Wesleyan University, A new approach for reconstructing

paleoclimatic and paleoecological variables from the sizes and shapes of fossil leaves.Richard Schulhof, Harvard University–Arnold Arboretum, Management response to hemlock woolly adelgid.Conghe Song, University of North Carolina and Harvard Bullard Fellow, Impacts of landuse history on regional

carbon budget: a remote sensing approach.Conrad Vispo, Hawthorne Valley Farm, New York, A preliminary description of the history and ecology of an eastern

New York farmscape.John Weishampel, University of Central Florida and Harvard Bullard Fellow, Exploring forest textures.Guthrie Zimmerma, University of Minnesota, Ruffed grouse habitat use in Minnesota: integrating multiple spatial

scales for management.

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40

Logs produced from the logging manipulation plots in the Hemlock Removal Experiment are loaded onto a log truck.Some of this wood was sawn and used for the new garage at the Forest.

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FOREST MANAGEMENT ANDINFRASTRUCTURE

Several significant projects were undertaken duringthe past two years, all with long-term impacts on thefuture of research and education at the Forest.

Shaler Hall underwent its first extensive renova-tion since it was built in 1942, with the assistance ofKuhn Riddle Architects and Teagno Construction,Inc. from Amherst. The first- and second-floor officesand meeting spaces were renovated with new light-ing, ceilings, doors, electrical systems, flooring andpaint. In addition to having an attractive new appear-ance, the space is now more energy efficient, providesnoise control, creates additional universally accessi-ble meeting and classroom spaces and providesincreased life safety protections with a new fire alarmsystem. The common room area was completely ren-ovated and several walls were removed to create aflexible education and research space for groups.With a full multimedia AV suite, classes ranging fromthree to fifty can comfortably meet for formal pre-sentations or informal meals.

In addition, the woods crew has been busy com-pleting a new maintenance garage on Prospect HillRoad, a quarter of a mile beyond the Fisher House.The new two-bay garage will reduce the impact ofForest operations on research sites and the watershedthat serves Prospect Hill. The garage, which is 30' ×52', was constructed of lumber that was harvested,milled, and planed on site by the woods crew. Thegarage features a composting toilet, a solid fuel heat-ing system, and will have an array of solar panels toproduce electricity on site. All work except for thefoundation and electricity was performed by the Harvard Forest woods crew. The total cost of thebuilding is estimated to be $72,000 exclusive of Forest staff time.

An ancillary project of the garage constructionwill be the installation of a new solar array with amaximum rated output of 9.98kW (DC), providingbetween 10–15 percent of the electricity needed by allof Prospect Hill research projects (EMS Tower, Hem-lock Walk Up, Little Prospect Hill, and Soil Warm-ing) as well as the new garage. The solar array willcost an estimated $95,000, with approximately$30,000 being provided by the Massachusetts Tech-nology Collaborative Small Renewables Initiativeand $20,000 by a matching grant from the FASDean’s Office. The garage will also be serviced by anew electric line from the main Prospect Hill primary

connection. This line will provide single-phase powerto the garage and triple-phase power to the sawmill.Eventually, this line will be connected up with theelectrical line running on Locust Opening Road, cre-ating a complete electrical circuit to the experimentson Prospect Hill.

Other significant projects included renovating aTorrey ecophysiology lab for general-use plant labspace. Two Community House apartments were con-verted to use for long-term dorm stays. Finally, thecrew assisted with rebuilding the soil warming shedafter it burned after a lightening strike in July 2005.

LIBRARY AND ARCHIVES

Judy Warnement and Sheila Connor from the Botan-ical and Arnold Arboretum Libraries continued tocatalog the library holdings in our Library of Con-gress classification. To date, 1,239 books have beenentered in the Harvard University (HOLLIS) cata-log. John Burk provided assistance to this project andalso completed sorting of the collections. During theprocess, material was transferred or donated to theHarvard Cabot Science and Wolbach Astrophysicslibraries, the Petersham Historical Society, the Barre,Royalston and Fitchburg public libraries, and theNational Weather Service. The library journals were

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sorted, inventoried (all foreign titles were transferredto the Cabot Library in February 2005), and alpha-betized. With input from Judy, Sheila, and other Har-vard Forest staff, the library committee of EmeryBoose, John Burk, Edythe Ellin, and Glenn Motzkincontinued meetings to plan future library projects,including a collections development policy.

As preliminary work for the LDI (Library Digi-tal Initiative) project, John Burk, Julie Pallant, andsummer proctor Hilary Crowell expanded andupdated the archive databases, and inventoried sev-eral of the collections that were previously uncata-loged, including the Forest’s large early twentieth-century historical photograph collection.

In conjunction with the land-management planand various administrative projects, the Harvard For-est property, deed, and map files were expanded andcataloged. A new flatbed scanner, with photo restora-tion software, was purchased and used regularly fora variety of projects.

INFORMATION MANAGEMENT ANDTECHNOLOGY ADVANCEMENT

Research data and documentation from studies con-ducted at the Forest are available in the DataArchive(http://harvardforest.fas.harvard.edu/data/archive.html) on the Harvard Forest web page. TheData Archive was extensively revised and updatedthis year in conjunction with submission of theLTER IV renewal proposal.

Harvard Forest datasets are now registered in adatabase at the LTER Network Office (http://metacat.lternet.edu) where they may be searched alongwith datasets from other LTER sites.

The Schoolyard LTER web page (http://harvardforest.fas.harvard.edu/museum/schoolyard.html) wasalso completely revised and updated this year to in-clude field data collected by participating K-12 classes.

ACTIVITIES OF THE HARVARD FOREST STAFF

Audrey Barker Plotkin coordinated the annual Har-vard Forest Ecology Symposium and Friday seminarseries at the Forest. She mentored three undergradu-ate students and served as a member of MathewTrumbull’s undergraduate capstone project commit-tee at Hampshire College.

Emery Boose served as a reader for a senior thesis at Brown University on hurricane impacts in

Taiwan. He gave talks on hurricane studies at theUniversity of Massachusetts, and at the annual Eco-logical Society of America meeting in Montreal.Emery also attended annual meetings of the LTERInformation Managers Committee, the LTER Infor-mation Managers Executive Committee, and LTERNetwork Information System Advisory Committee.

John Burk continued to attend and participate inconservation groups, including the Tully Lake andQuabbin use meetings, and led several local fieldtrips. John contributed to several publications, includ-ing the Mount Grace Conservation Trust twentiethanniversary book.

Betsy Colburn was the keynote speaker at theCleveland Museum of Natural History’s conserva-tion symposium: “Ephemeral Gems: Exploring Sea-sonal Ponds,” and she presented public lectures onvernal pool ecology and the application of the “Wild-lands and Woodlands” approach to pool conserva-tion at the Lincoln (Massachusetts) Garden Club, theLowell National Historic Park, and the NorthwestConservation District of Connecticut. Betsy coau-thored the chapter on invertebrates in a new book,focusing on best management practices for activitiescarried out in the vicinity on vernal pools, and wasalso asked to speak on the importance of funding forsystematics and natural history research at the specialsession on systematics and life history at the NorthAmerican Benthological Society Annual Meeting inAnchorage, Alaska in June 2006.

Aaron Ellison spent a modest amount of time onthe road, riding the rails, and in the air this year. Inaddition to his apparently annual lecture on Bayesianstatistics at the University of Massachusetts, Aarongave a keynote address at the second Mangrove Macrobenthos Meeting in Australia. This meeting

REU student Mathew Trumbull.

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afforded him the opportunity to revisit research areasand data that he has not paid much attention to fornearly a decade. Aaron also visited Howard Univer-sity and Delaware State University, where he pre-sented research seminars and recruited students forthe Harvard Forest summer research program inecology, and he was invited to a four-day workshopon uncertainty in ecological analysis at the Mathe-matical Biosciences Institute at Ohio State, where hepresented a talk on statistical challenges in detectingsudden ecosystem changes and alternative stablestates in ecological communities. Aaron was appoint-ed an Adjunct Professor in the Biology Departmentat the University of Massachusetts, and is currentlysupervising his first UMass Ph.D. student, SydneRecord. He is also on the M.Sc. committee of JustineRoths at the University of Southern Maine at Port-land, who is testing a hypothesis that Aaron pro-posed in 1991, that moths can alter competitivedynamics between two salt-marsh plants. Aaron ishosting a post-doc, Jim Karagatzides (Queen’s Uni-versity, Ontario), who received a two-year fellowshipfrom NSERC, Canada’s NSF-equivalent. Aaroncontinued his stints as associate editor-in-chief forEcology and Ecological Monographs, as a member of theEcological Society’s publications committee, and as asubject matter editor for Ecology Letters. Closer tohome, Aaron continued his terms on the RoyalstonConservation Commission and the Royalston OpenSpace Committee, and stepped down from the LandProtection Committee of the Mt. Grace Land Trustafter three years of work. Perhaps most importantly,Aaron’s ongoing work with the Friends of Tully Lakeled to a key land-use victory in June 2006 when theAthol Planning Board rejected a definitive plan for asubdivision to be built overlooking Tully Lake.

In September, Brian DeGasperis attended anInvasive Plant Summit in Framingham, Massachu-setts, evaluating the interactions between historicalland use and modern disturbances.

Ed Faison spent the summer and early fall of2005 in the Quabbin and Ware River forests gather-ing data for his MFS thesis on moose-foraging ecolo-gy. He also helped initiate a pilot study of moosemovements in Royalston, Massachusetts for themonth of July and advised Christian Foster and BenEwing for this project. During the fall and winter, Edcollaborated with David Foster, Wyatt Oswald, andElaine Doughty on three paleoecology papers. InMarch of 2006, Ed gave a talk at the Athol Bird andNature Club on moose, and in April was awardedHarvard University’s Bowdoin Prize for his essay onthe paleoecology of ragweed. In May Ed successfullydefended his thesis and began a research assistantposition at Highstead Arboretum in Redding, Con-necticut where he collaborates with David Foster ona study of invasive species and other vegetation pat-terns. Ed advises two REU students Dana Graef andLisa Schauer for this project at Highstead. In June Edattended Harvard’s graduation along with MFSclassmate Brooks Mathewson.

Brian Hall acted as reviewer for articles pub-lished in Landscape Ecology, Journal of Biogeography, andNortheast Naturalist. Brian also continued with onlinecourses in Visual Basic for Applications for program-ming in GIS software for automating repetitive tasks.

Jim Karagatzides is coinvestigator for the “YouthScience Outreach in the Mushkegowuk Territory ofSub-Arctic Ontario, Canada.” Jim’s contributionincludes field trips to the local muskeg using theJessica Butler with pitcher plants in the greenhouse.

Ed Faison conducting vegetation surveys.

northern pitcher plant (Sarracenia purpurea) as a micro-ecosystem to teach food web dynamics. In this con-text of understanding a changing environment, theyinclude bird surveys and fossil hunts in the region.

Dave Kittredge gave several local presentationson the wildlands and woodlands vision to land trustsand watershed associations. He has sought examplesof woodland council functionality as part of thisgreater wildlands and woodlands vision. He contin-ued his collaboration on analysis of the forest harvestdata (McDonald, et al. 2006). Dave also published apaper in the Natural Areas Journal on timber har-vesting and the protection of rare species habitat inMassachusetts.

Heidi Lux worked on HWA projects at theArnold Arboretum, the Simes Tract of Harvard For-est, and nine ecosystem sites in Connecticut. Heidiattended the ninetieth ESA annual meeting in Mon-treal, Canada in August of 2005, presenting HWAresearch at a poster session titled: “Changes inEcosystem Response Associated with Hemlock Wool-ly Adelgid Infestation and Preemptive Logging inSouthern New England.” In January of 2006 Heidipresented on the current status of Harvard Forestresearch on Hemlock Hill at the Arnold Arboretum inJamaica Plain, Boston: “Hemlock Woolly Adelgid andLogging at the Arnold Arboretum: Pre- and Post-treatment Results.” At the Harvard Forest Symposiumin March of 2006, Heidi gave a talk titled: “Ecosys-tem Responses to Forest Cutting after HWA Infesta-tion – New Insights from an Urban Site.” In the sum-mer of 2006, Heidi assisted David Orwig with thementoring of REU student Christy Rollinson. High-school student Eowyn Connolly-Brown continuedwith the HWA group through the early summer of2006, with Heidi as her main supervisor.

Robert McDonald spent much of his year writinga manuscript in conjunction with the Agricultural

Landscapes in Transition project. This manuscriptinvolves collaboration among many personnel atHarvard Forest, Coweeta Hydrological Lab, andPhoenix/Central Arizona LTERs. Rob has presentedarticles at two conferences, one in San Diego at themeeting of the International Association of Land-scape Ecologists and one in San Jose at the meetingof the Society for Conservation Biology.

Since July 1, 2005, Jackie Mohan and the groupfrom MBL (Marine Biological Laboratory) has started a new physiological girdling study of foresttrees and started a new study of fine-root biomassresponses to warming. They have also continuedtheir extensive research work (biogeochemistry,growth, survivorship, reproduction, tissue chemistry,litter production, and decomposition) at their two soilwarming experiments. In summer 2006 Jackie(MBL) mentored REU student, Rebecca Orozco.Jackie has two new RAs: Rob Hanifin and KatherineLenoir (Rob was REU student in the summer of2004). Jackie also gave seminars at Purdue Univer-sity, Michigan State University, the Kellogg Biolog-ical Station at Indiana University, and the Universityof Texas–Austin, during the spring of 2006.

Glenn Motzkin continues to serve as an ecologyadvisor for The Trustees of Reservations, an associ-ate member of the Massachusetts Natural Heritageand Endangered Species Program Advisory Com-mittee, and a member of the board of editors forNortheastern Naturalist. Glenn also served on the thesiscommittees of Posy Busby, Gretel Clarke (UMass),Brian DeGasperis, and Ed Faison and led a field tripto the Connecticut River for an ENPP class in Con-servation Biology and a trip to Montague Plain forthe Conway School of Landscape Design. Glenn,

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Rob McDonald.

Jackie Mohan, Rob Hanifin and Liz Burrows.

Dave Orwig, and John O’Keefe led field trips for theFreshman Seminar class.

In 2004-05 John O'Keefe gave talks on HarvardForest research and the history of northeasternforests at the annual meeting of the MassachusettsChapter of the American Chestnut Foundation, theMillers River Environmental Center, and the Townof Bedford Arbor Day Commemoration and in2005–06 at the Pioneer Valley Institute in October,the Massachusetts Audubon Society’s ArcadiaNature Center in March and the Eastern ForestResource Program Leaders Conference in May. In2005 John completed nine years (the maximum con-secutive years allowed) as a director and vice presi-dent of Mount Grace Land Conservation Trust(MGLCT) and stepped down. He continues to serveon the MGLCT Land Committee, on the boards ofthe Massachusetts Forestry Association, MillersRiver Environmental Center, and the SpannocchiaFoundation, on the executive committee of the NorthQuabbin Regional Landscape Partnership, and onthe steering committee of the Quabbin to Cardigan(Q2C) conservation collaborative. He also continuesto serve on the secretary of Environmental Affairs’Advisory Group on Environmental Education. InMarch 2005, John and his wife, Lynne Stopen, trav-eled to China to meet and bring home their seconddaughter, Erin.

Dave Orwig gave invited talks about hemlockwoolly adelgid at Williams College in Williamstown,Massachusetts, and at Westfield State University inWestfield, Massachusetts. With David Kittredge, hespoke about managing hemlock forests at the South-ern New England Logger Education Program inGardner, Massachusetts. He led field trips at HarvardForest, Ashburnham Elementary School, and War-wick Elementary School as part of the HF LTERSchoolyard Program and as part of the Harvard Uni-versity Freshman Seminar Class. Dave continued toserve on the Ph.D. committee of Kenn Clark and asthe Ph.D. dissertation advisor to Anthony D’Amatoat the University of Massachusetts. Dave activelyparticipated in a working group on Pests, Pathogens,and Invasive species held at UMass, Boston as partof a larger effort run by the coalition, NE Forests2100.

Julie Pallant attended an ArcGIS training sessionto become ESRI certified in November 2005, spon-sored by the LTER Network Office.

P. Barry Tomlinson began a survey of the anato-my of all palms in Miami at Fairchild Tropical Botan-

ical Garden, Coral Gables, Florida, October 2005 toApril 2006. Emphasis is on leaf anatomy and usesextensive collections of palms cultivated in Floridaand Hawaii. Barry’s visit to New Zealand in Febru-ary–March 2006 in the company of Professor FrancisHallé of Montpellier, France, in order to study archi-tecture of native and introduced trees demonstratedthe exceptional diversity of tree form concomitantwith the vegetational history of New Zealand and itsstrong tropical components.

LAND PROTECTION EFFORTS

Previous reports have outlined the increasing threatto Harvard Forest’s land base, research, and educa-

tional activities posed by the extensive exposure of itsland to abutting public roads and developable privateproperty. Fueled by rapid increases in housing pricesand densities in areas to our east and improved trans-portation, our region is experiencing major increasesin land prices and development. This developmentthreatens Harvard Forest’s hundred-year record ofexperiments and studies that gain irreplaceable valuethrough time, many involving sensitive organisms orecosystem processes such as gas fluxes and hydrolo-gy that are extremely vulnerable to even the indirectimpacts of human activity. In response to thesethreats Harvard Forest has initiated major efforts to

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REU Student Lori Simper.

work with local government, regional and statewideconservation groups, state agencies, and the federalgovernment to establish a buffer of protected landand ensure the Forest’s continued viability as aresearch and educational facility.

The region surrounding the northeast corner ofthe Quabbin Reservation, which encompasses Har-vard Forest, has been identified as an important focusarea for land conservation by collaborative groupsworking at several scales. It has been selected as oneof six initial focus areas by the Quabbin to CardiganConservation Collaborative (Q2C), a group of pub-lic and private conservation stakeholders led by theSociety for the Protection of New Hampshire Forests(SPNHF).

A Harvard Forest study published in 1997 (Fos-ter and Golodetz) documented a haphazard patternof land protection within the north Quabbin region.This report led to the creation of the North QuabbinRegional Landscape Partnership (NQRLP) to coor-dinate all the public and private groups interested inland protection within a twenty-six town region.

In the fall of 2005, under the direction of JohnO’Keefe, Harvard Forest convened the East QuabbinLand Protection Group. With representatives fromlocal towns (Athol, Barre, Hardwick, Petersham,Phillipston), regional land trusts, and nonprofits(Mount Grace Land Conservation Trust, East Quab-bin Land Trust, The Trustees of Reservations, Mass-achusetts Audubon Society) and state agencies (DCRand DFG) this group seeks to identify priorities andcoordinate land-protection activities within ourimmediate region. One outcome has been the verywell supported (and ranked third nationally in FY06)U.S. Forest Service Forest Legacy program “QuabbinCorridor” grant application requesting funds to pur-chase conservation easements covering more than2,000 acres on twenty-one properties in Petersham,Phillipston, and Barre. We anticipate sufficient fund-ing to establish easements on two parcels abuttingour Prospect Hill tract this year, the hundred-acreWilson property abutting the north boundary andthe sixty-eight-acre Bryant woodlot abutting thesouth boundary. Once these restrictions are in place,Harvard Forest will purchase the restricted propertiesas part of its buffer-creation plan. These purchaseswill be possible thanks to the very generous dona-tions and pledges from our friends, alumni, neigh-bors, regional groups and foundations in support ofour land-protection efforts. These two parcels repre-sent the start of a strategic plan to provide a buffer of

protected land around our research and educationaloperations.

In a related planning effort, John O’Keefe andAudrey Barker Plotkin are working with HarvardForest staff and other senior researchers to develop amaster plan for the Forest’s properties. This plan willconsolidate historical data for all the properties anddefine and map zones within Harvard Forest thatwould be appropriate for different levels of research,educational, recreational, and management activity.Certain zones would be restricted to minimal impactuses, other zones would be available for generalresearch with consideration for any impacts to past orongoing research, and other limited zones would bezoned for future infrastructure expansion if the needshould arise. This master plan will provide a docu-ment to guide the use and management of HarvardForest’s properties as the Forest prepares to enter itssecond century.

46

John Aber University of New Hampshire

Jacqueline Aitkenhead University of New Hampshire

Kate Ackerman U.S. Geological SurveyLynn Adler University of MassachusettsJeanne Anderson University of New

HampshireMark Ashton Yale UniversityPaul Barten University of MassachusettsMark Battle Bowdoin CollegeBryan Blair NASA Goodard Space Flight

CenterJoe Blanchard Ecosystems Center — MBLRichard Bowden Allegheny College Frank Bowles Ecosystems Center — MBLHilary Callahan Barnard College, Columbia

UniversityCynthia Chang University of MarylandJonathan Chen Oberlin CollegeElizabeth Chilton University of Massachusetts

Bridget Collins Holy Cross CollegeScott Costa University of VermontPatrick Crill University of New

HampshireBryan Dail University of MaineMichael Daley Boston UniversityTony D’Amato University of MassachusettsEric Davidson Woods Hole Research

CenterKatrina del Fierro Barnard College, Columbia

UniversityR. Drijber University of NebraskaRalph Dubayah University of MarylandLuke Durbin Illinois Wesleyan UniversityGeoffrey Dutton University of Colorado and

NOAA/CMDLJim Ehleringer University of UtahJames Elkins University of Colorado and

NOAA/CMDLSerita Frey University of New

Hampshire

47

VISITING RESEARCH SCIENTISTS AT THE HARVARD FOREST 2004–2005

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

Heidi Fuchs Amherst CollegeDavid Geiser Pennsylvania State

UniversityNicholas Gotelli University of VermontKelly Grogan Dartmouth CollegeElizabeth Hane Rochester Institute of

TechnologyErik Hobbie University of California,

BerkeleyMichelle Hofton University of MarylandDavid Hollinger USDA Forest ServiceHolly Hughes Woods Hole Research

CenterPeter Hyde University of MarylandScott Isard Pennsylvania State

UniversityGordon Jacoby Columbia UniversityJerry Jenkins White Creek Field SchoolChelsea Kammerer- Clark University

BurnhamJim Karagatzides Queen’s University, OntarioSylvan Kaufman Atkins ArboretumJoel Kingsolver University of North

CarolinaJohn Klironomos University of Guelph,

OntarioRobert Knox NASA Goddard Space

Flight CenterDöerte Köester University of Waterloo,

OntarioEleanor Lahr Ithaca CollegeChun-Ta Lai University of UtahMatthew Lau Humboldt State UniversityJohn Lee University of MaineXuhui Lee Yale UniversityWendy Liu University of California–

BerkeleyAnna Mari Lone Amherst CollegeStephanie Long USDA Forest ServiceFrank Lowenstein The Nature ConservancyAlison Magill University of New

HampshireMegan Manner Duke UniversityLynn Margulis University of MassachusettsMary Martin University of New

HampshireCharles McClaugherty Mount Union CollegeKarena McKinney Amherst CollegeKirsten McKnight Brigham Young University Mitch Mulholland University of MassachusettsJerry Melillo Ecosystems Center — MBL

Qilong Min SUNY–AlbanyRakesh Minocha USDA Forest ServiceJacqueline Mohan Ecosystems Center — MBLThomas Muller Centre for Agricultural

Lanscape and Land UseResearch–Muncheberg

Knute Nadelhoffer University of MichiganDonald Niebyl Great Basin InstituteScott Ollinger University of New

HampshireColin Orians Tufts UniversityAngelica Patterson Barnard College, Columbia

UniversityNeil Pederson Columbia UniversityBrigit Peterson University of MarylandCory Pettijohn Boston UniversityNathan Phillips Boston UniversityNick Povak University of Wisconsin–

MadisonGreg Ragland University of North

CarolinaAndrew Richardson University of New

HampshireChuck Rodrigues University of MaineNicholas Rosenstock University of California–

BerkeleyScott Saleska University of ArizonaKathleen Savage Woods Hole Research

CenterNeal Scott Woods Hole Research

CenterTristram Seidler Imperial College at Silwood

Park, UKTim Sipe Franklin & Marshall CollegeHeather Smith University of New

HampshireMarie-Louise Smith USDA Forest ServiceWilliam Sobczak Holy Cross CollegeBernhard Stadler Bayreuth Institute for

Terrestrial EcosystemResearch

Paul Steudler Marine Biological Laboratories

Eric T Sundquist U.S. Geological SurveyRuth K Varner University of New

HampshireRebecca Waysek U.S. Geological SurveyHui-Ju Wu Yale UniversityHina Zafar Barnard College, Columbia

University

48

John Aber University of New Hampshire

Jacqueline Aitkenhead University of New Hampshire

Kate Ackerman U.S. Geological SurveyLynn Adler University of MassachusettsMark Ashton Yale UniversityPaul Barten University of MassachusettsMark Battle Bowdoin CollegePeter Bettmann- Hampshire College

KersonJoe Blanchard Ecosystems Center — MBL

Richard Bowden Allegheny College Frank Bowles Ecosystems Center — MBLAlfram Bright Harvard UniversityTimothy Brodribb Harvard UniversityCaroline Brophy University College, DublinRobert Buchsbaum Massachusetts Audubon

SocietyJohn Budney Harvard UniversityHilary Callahan Barnard College, Columbia

UniversityAndrew Cavanagh University of MassachusettsCynthia Chang University of MarylandJonathan Chen Oberlin CollegeElizabeth Chilton University of MassachusettsBrendan Choat Harvard UniversityAimee Classen Oak Ridge National

LaboratoryBridget Collins Holy Cross CollegeAmanda Cooper University of Central

FloridaScott Costa University of VermontPatrick Crill University of New

HampshireDaniel Curran Harvard UniversityBryan Dail University of MaineMichael Daley Boston UniversityEric Davidson Woods Hole Research

CenterDavid Diaz Harvard UniversityKatrina del Fierro Barnard College, Columbia

UniversityKathleen Donohue Harvard UniversityR. Drijber University of NebraskaRalph Dubayah University of Maryland

David Dudley USDA — States Departmentof Agriculture

Luke Durbin Illinois Wesleyan UniversityGeoffrey Dutton University of Colorado and

NOAA/CMDLJim Ehleringer University of UtahJames Elkins University of Colorado and

NOAA/CMDLJoe Elkinton University of MassachusettsDavid Fitzgarrald State University of New

York–AlbanySerita Frey University of New

HampshireHeidi Fuchs Amherst CollegeDavid Geiser Pennsylvania State

UniversitySandy Gillespie University of MassachusettsNicholas Gotelli University of VermontKelly Grogan Dartmouth CollegeElizabeth Hane Rochester Institute of

TechnologyKristen Hladun University of MassachusettsErik Hobbie University of California–

BerkeleyMichelle Hofton University of MarylandMichele Holbrook Harvard UniversityDavid Hollinger USDA Forest ServiceHolly Hughes Woods Hole Research

CenterPeter Hyde University of MarylandScott Isard Pennsylvania State

UniversityGordon Jacoby Columbia UniversityJerry Jenkins White Creek Field SchoolChristine Jones Harvard UniversityChelsea Kammerer- Clark University

BurnhamJim Karagatzides Queen’s University, OntarioJennifer Karberg Michigan Technological

UniversitySylvan Kaufman Atkins ArboretumGeeta Kharkwal Boston CollegeJoel Kingsolver University of North

CarolinaJohn Klironomos University of Guelph,

Ontario

49

VISITING RESEARCH SCIENTISTS AT THE HARVARD FOREST 2005–2006

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

Robert Knox NASA Goddard Space Flight Center

Döerte Köester University of Waterloo,Ontario

Eleanor Lahr Ithaca CollegeChun-Ta Lai University of UtahMatthew Lau Humboldt State UniversityJohn Lee University of MaineXuhui Lee Yale UniversityWendy Liu University of California

BerkeleyAnna Mari Lone Amherst CollegeStephanie Long USDA Forest ServiceFrank Lowenstein The Nature ConservancyAlison Magill University of New

HampshireMegan Manner Duke UniversityLynn Margulis University of MassachusettsMary Martin University of New

HampshireCharles McClaugherty Mount Union CollegeKathryn McKain Harvard UniversityKarena McKinney Amherst CollegeKirsten McKnight Brigham Young UniversityJason McLachlan University of California–

DavisMitch Mulholland University of MassachusettsJerry Melillo Ecosystems Center — MBLQilong Min SUNY–AlbanyRakesh Minocha USDA Forest ServicePaul Moorcroft Harvard UniversityMichael Muilenberg Harvard School of Public

HealthWilliam Munger Harvard UniversityThomas Muller Centre for Agricultural

Lanscape and Land UseResearch–Muncheberg

Knute Nadelhoffer University of MichiganHoward Neufeld Appalachian State

UniversityDonald Niebyl Great Basin InstituteWenge Ni-Meister Hunter College–CUNYScott Ollinger University of New

HampshireColin Orians Tufts UniversityRobert Packard University of MassachusettsAnnie Paradis University of MassachusettsAngelica Patterson Barnard College, Columbia

UniversityNeil Pederson Columbia UniversityBrigit Peterson University of Maryland

Celeste Peterson Harvard UniversityCory Pettijohn Boston UniversityNathan Phillips Boston UniversityNick Povak University of Wisconsin–

MadisonAnne Pringle Harvard UniversityElizabeth Hammond Harvard University

PyleGreg Ragland University of North

CarolinaPeter Raymond Yale University Sydne Record University of Massachusetts Andrew Richardson University of New

HampshireChuck Rodrigues University of MaineChristine Rogers Harvard UniversityNicholas Rosenstock University of California–

BerkeleyScott Saleska University of ArizonaNate Sanders University of Tennessee Kathleen Savage Woods Hole Research

CenterSanna Sevanto Harvard UniversityNeal Scott Woods Hole Research

CenterTim Sipe Franklin & Marshall CollegeHeather Smith University of New

HampshireMarie-Louise Smith USDA Forest ServiceSeeta Sistla Brown UniversityWilliam Sobczak Holy Cross CollegeBernhard Stadler Bayreuth Institute for

Terrestrial EcosystemResearch

Paul Steudler Marine BiologicalLaboratories

Eric Sundquist U.S. Geological SurveyPamela Templer Boston UniversitySusan Trumbore University of CaliforniaRuth Varner University of New

HampshireRebecca Waysek U.S. Geological SurveyJohn Weishampel University of Central

FloridaSteve Wofsy Harvard UniversityBenjamin Wolfe Harvard UniversityHui-Ju Wu Yale UniversityHina Zafar Barnard College, Columbia

University

50

PUBLICATIONS

*Indicates Summer Research Program forUndergraduates participant

Aber, J. D. and A. H. Magill. 2004. Chronic nitrogenadditions at the Harvard Forest: The first fifteenyears of a nitrogen saturation experiment. ForestEcology and Management 196:1-5.

Atwater*, D. Z., J. L. Butler, and A. M. Ellison. 2006.Spatial distribution and impacts of moth her-bivory on northern pitcher plants. NortheasternNaturalist 13:43–56.

Baillie, I. C., P. S. Ashton, S. P. Chin, S. J. Davies, P.A. Palmioto, S. E. Russo, and S. Tan. 2006. Spa-tial associations of humus, nutrients and soils inmixed dipterocarp forest at Lambir, Sarawak,Malaysian Borneo. Journal of Tropical Ecology22:543–553.

Bain*, W. G., L. Hutyra, D. C. Patterson*, A. V.Bright, B. C. Daube, J. W. Munger, and S. C.Wofsy. 2005. Wind-induced error in the meas-urement of soil respiration using closed dynamicchambers. Agricultural and Forest Meteorology131:225–232.

Bakwin, P. S., K. J. Davis, C. Yi, S. C. Wofsy, J. W.Munger, L. Haszpra, and Z. Barcza. 2004.Regional carbon dioxide fluxes from mixing ratiodata. Tellus 56B:301–11.

Bank, M. S., C. S. Lotfin, and E. R. Jung. 2005. Mer-cury bioaccumulations in Northern two-linedsalamanders from streams of the NortheasternU.S. Excotoxicology 14:181–191.

Bauer, G. A., F. A. Bazzaz, R. Minocha, S. Long, A.Magill, J. Aber, and G. M. Berntson. 2004.Effects of chronic N additions on tissue chem-istry, photosynthetic capacity, and carbon seques-tration potential of a red pine (Pinus resinosa Ait.)stand in the N. E. United States. Forest Ecology andManagement 196:173–186.

Belin, D. L., D. B. Kittredge, T. H. Stevens, D. Den-nis, C. M. Schweik, and B. J. Morzuch. 2005.Assessing private forest owner attitudes towardecosystem-based management. Journal of Forestry103(1):28–33.

Bellemare, J., G. Motzkin, and D. R. Foster. 2005.Rich mesic forests: edaphic and physiographicdrivers of community variation in western Mass-achusetts. Rhodora 107:239–283.

Bertin, R. I., B. G. DeGasperis, and J. M. Sabloff.2006. Land use and forest history in an urban

sanctuary in central Massachusetts. Rhodora108:119–141.

Boose, E. R. 2004. A method for reconstructing his-torical hurricanes. Pp. 99–120 in Hurricanes andTyphoons: Past, Present, and Future. R. Murnane andK. Liu (Eds.). Columbia University Press, NewYork.

Borken W., K. Savage, E. A. Davidson, and S. E.Trumbore. 2006. Effects of experimental droughton soil respiration and radiocarbon efflux from atemperate forest soil. Global Change Biology12:177–193.

Bowden, R. D., E. Davidson, K. Savage, C. Arabia*,and P. Steudler. 2004. Chronic nitrogen addi-tions reduce total soil respiration and microbialrespiration in temperate forest soils at the Har-vard Forest. Forest Ecology and Management 196:43–56.

Burton, P. J. 2005. Ecosystem management and con-servation biology. Pp. 307–322 In S.B. Watts andL. Tolland (eds.), Forestry Handbook for BritishColumbia. Faculty of Forestry, University ofBritish Columbia, Vancouver, B.C.

Butler, J., D. Atwater*, and A. Ellison. 2005. Red-spotted newts: An unusual nutrient source fornorthern pitcher plants. Northeastern Naturalist12:1–10.

Chan, A. S. K., P. A. Steudler, R. D. Bowden, J.Gulledge, and C. Cavanaugh. 2005. Conse-quences of nitrogen fertilization on soil methaneconsumption in a productive temperate decidu-ous forest. Biology and Fertility of Soils 41:182–189.

Chapman, M. 2004. The Federal role in advancinglandscape-scale, multi-owner forest conservationand stewardship. Pp. 51–64 in Forest Conservationand Stewardship in Massachusetts. C. H. W. Fosterand P. R. Hagenstein (Eds.). Harvard Forest,Petersham, Massachusetts.

Choat, B., E. Lahr*, P. J. Melcher, M. A. Zwieniecki,and N. M. Holbrook. 2005. The spatial patternof air seeding thresholds in mature sugar mapletrees. Plant, Cell and Environment 28:1082–1089.

Clark, S. A. 2004. Stewards of our land: the forestservice’s stewardship contracting program andthe management of public land in regard to wild-fires. B.A. Thesis, Harvard College.

Cobb, R. C., D. A. Orwig, and S. Currie*. 2006.Decomposition of green foliage in eastern hem-lock forests of southern New England impactedby hemlock woolly adelgid infestations. CanadianJournal Forest Research 36:1–11.

51

Colburn, E. A. 2004. Vernal Pools: Natural History andConservation. McDonald & Woodward, Granville,OH.

Compton, J. E., L. S. Watrud, L. A. Porteous, and S.DeGrood*. 2004. Response of soil microbialmiomass and community composition to chronicnitrogen additions at Harvard Forest. Forest Eco-logy and Management 196:143–158.

Currie, W. S., K. J. Nadelhoffer, and J. D. Aber. 2004.Redistributions of 15N highlight turnover andreplishment of mineral soil organic N as a long-term control on forest C balance. Forest Ecologyand Management 196:109–127.

Davidson, E. A. and I. A. Janssens. 2006. Tempera-ture sensitivity of soil carbon decomposition andfeedbacks to climate change. Nature 440:165–173.

Davidson, E. A., I. A. Janssens, and Y. Luo. 2005.On the variability of respiration in terrestrialecosystems: moving beyond Q10. Global ChangeBiology 12:154–164.

Davidson, E. A., K. Savage, S. E. Trumbore, and W.Borken. 2006. Vertical partitioning of CO2 pro-duction within a temperate forest soil. GlobalChange Biology 12:944–956.

DeBell, L. J., R. W. Talbot, J. E. Dibb, J. W. Munger,E. V. Fischer, and S. E. Frolking. 2005. A majorregional air pollution event in the northeasternU.S. caused by extensive forest fires in Quebec,Canada. Journal of Geophysical Research-Atmospheres109.

Dixon, P. M., A. M. Ellison, and N. J. Gotelli. 2005.Improving the precision of estimates of the fre-quency of rare events. Ecology 86:1114–1123.

Dixon, P. M., A. M. Ellison, and N. J. Gotelli. 2005.Improving the precision of estimates of the fre-quency of rare events. Ecology 86:1114–1123.

Donohue, K. 2004. Density-dependent multilevelselection in the Great Lakes sea rocket. Ecology85:180–191.

———. 2005. Niche construction through phenologicalplasticity: Life history dynamics and ecologicalconsequences. (Invited review) New Phytologist166:83–92.

———. 2005. Seeds and seasons: Interpreting germina-tion timing in the field. Seed Science Research15:175–187.

Donohue, K., C. G. L. Dorn, E. S. Kim, A. Aguilera,C. R. Polisetty, and J. Schmitt. 2005. Environ-mental and genetic influences on the germinationof Arabidopsis thaliana in the field. Evolution59:740–757.

———. 2005. The evolutionary ecology of seed germi-nation of Arabidopsis thaliana: Variable naturalselection on germination timing. Evolution59:758–770.

———. 2005. Niche construction through germinationcueing: Life history responses to timing of germi-nation in Arabidopsis thaliana. Evolution59:771–785.

Donohue, K., C. R. Polisetty, and N. J. Wender*.2005. Genetic consequences of niche construc-tion: Plasticity-mediated genetic constraints onthe evolution of seed dispersal of Arabidopsisthalinia (Brassicaceae). The American Naturalist165:537–550.

Douglas, T. and P. J. Burton. 2005. Integratingecosystem restoration into forest management inBritish Columbia, Canada. Pp. 423–444 in J. A.Stanturf and P. Madsen (eds.), Restoration of Bore-al and Temperate Forests. CRC Press. Boca Raton,Florida.

Ellison, A. M. 2004. Commentary on statistics andthe scientific method in ecology by Brian Dennis.Pp. 362–367 in The Nature of Scientific Evidence. M.L. Taper and S. R. Lele (Eds.). University ofChicago Press, Chicago, Illinois.

———. Wetlands of Central America. Wetlands Ecologyand Management 12:3–55.

———. Turning the tables: plants bite back. Wings25–30.

Ellison A. M., M. S. Bank, B. D. Clinton, E. A. Col-burn, K. Elliott, C. R. Ford, D. R. Foster, B. D.Kloeppel, J. D. Knoepp, G. M. Lovett, J. Mohan,D. A. Orwig, N. L. Rodenhouse, W. V. Sobczak,K. A. Stinson, J. K. Stone, C. M. Swan, J.Thompson, B. Von Holle, and J. R. Webster.2005. Loss of foundation species: consequencesfor the structure and dynamics of forestedecosystems. Frontiers in Ecology and the Environment479–486.

Ellison, A. M. and E. J. Farnsworth. 2005. The costof carnivory for Darlingtonia california (Sarracheni-aceae): Evidence from relationships among leaftraits. American Journal of Botany 92:1085–1093.

Ellison, A. M., H. L. Buckley, T. E. Miller, and N. J.Gotelli. 2004. Morphological variation in Sarrace-nia purpurea (Sarraceniaceae): geographic, envi-ronmental, and taxonomic correlates. AmericanJournal of Botany 91:930–1935.

Ellison, A. M., J. Chen*, D. Díaz*, C. Kammerer-Burnham*, and M. Lau*. 2005. Changes in antcommunity structure and composition associated

52

with hemlock decline in New England. In: ThirdSymposium on Hemlock Woolly Adelgid in the EasternUnited States.

Ellison, A. M., L. J. Osterweil, J. L. Hadley, A. Wise,E. Boose, L. Clarke, D. R. Foster, A. Hanson, D.Jensen, P. Kuzeja, E. Riseman, and H. Schultz.2006. Analytic Webs Support the Synthesis ofEcological Data Sets. Ecology 87:1345–1358.

Faison, E. K., D. R. Foster, W. W. Oswald, E. D.Doughty, and B. C. S. Hansen. 2006. Early-Holocene openlands in southern New England.Ecology 87:10:2537–2547.

Farnsworth, E. and S. Lowe. 2005. Peterson Field Guideto Ferns and Their Related Families. Houghton Mif-flin, Boston.

Farnsworth, E. J. and D. E. Ogurcak. 2006. Biogeog-raphy and decline of rare plants in New England:historical evidence and contemporary monitor-ing. Ecological Applications 16:1327–1337.

Finley*, A. O. and D. B. Kittredge. 2006. Thoreau,Muir, and Jane Doe: different types of privateforest owners need different kinds of forest man-agement. Northern Journal of Applied Forestry23:27–34.

Finley*, A. O., D. B. Kittredge, T. H. Stevens, C. M.Schweik, and D. Dennis. 2006. Interest in cross-boundary cooperation: identification of distincttypes of private forest owners. Forest Science52:10–22.

Foster, C. H. W., P. R. Hagenstein, and D. R. Foster.2004. Forest Conservation and Stewardship inMassachusetts. Pp. 1–10 in Forest Conservation andStewardship in Massachusetts. C. H. W. Foster and P.R. Hagenstein (Eds.). Harvard Forest, Peter-sham, MA.

Foster, D. R. 2006. The importance of land use his-tory to conservation biology. In M. J. Groom, G.K. Meffe, and C. R. Carroll (eds.), Principles ofConservation Biology. Sinauer Associates, Sunder-land, MA.

Foster, D. R., D. B. Kittredge, B. Donahue, G.Motzkin, D. A. Orwig, A. M. Ellison, B. Hall, B.Colburn, and A. D’Amato. 2005. Wildlands andWoodlands: A Vision for the Forests of Massachusetts.Harvard Forest Paper No. 27. Harvard Forest,Petersham, MA.

Foster. D. R. and D. A. Orwig. 2006. Pre-emptiveand salvage harvesting of New England forests:when doing nothing is a viable alternative. Con-servation Biology 20:959–970.

Francis, D. R. 2004. Distribution of midge remains

(Diptera: Chironomidae) in surficial lake sedi-ments in New England. Northeastern Naturalist11:459–478.

Frey, S. D. 2006. Forest ecosystem responses tochronic nitrogen additions. Teaching Issues andExperiments in Ecology 4.

Frey, S. D., M. Knorr, J. L. Parrent, and R. T. Simp-son. 2004. Chronic nitrogen enrichment affectsthe structure and function of the soil microbialcommunity in temperate hardwood and pineforests. Forest Ecology and Management196:159–171.

Gavin*, D. G., L. B. Brubaker, J. S. McLachlan, andW. W. Oswald. 2005. Correspondence of pollenassemblages with forest zones across steep envi-ronmental gradients, Olympic Peninsula, Wash-ington, USA. The Holocene 15:633–650.

Gotelli, N. J. and A. M. Ellison. 2004. A Primer of Eco-logical Statistics. Sinauer Associates, Sunderland,Massachusetts, U.S.A.

———. 2006. Food-web models predict species abun-dances in response to habitat change. PublicLibrary of Science Biology 14:e324.

———. 2006. Forecasting extinction risk with non-sta-tionary matrix models. Ecological Applications16:51–61.

Gu, L. H., E. M. Falge, T. Boden, D. D. Baldocchi,T. A. Black, S. R. Saleska, T. Suni, S. B. Verma,T. Vesala, S. C. Wofsy, and L. K. Xu. 2005.Objective threshold determination for nighttimeeddy flux filtering. Agricultural and Forest Meteorol-ogy 128:179–197.

Harper, K. A., S. E. Macdonald, P. J. Burton, J.Chen, K. D. Brosofske, S. C. Saunders, E. S.Euskirchen, D. Roberts, M. S. Jaiteh, and P-A.Esseen. 2005. Edge influence on forest structureand composition in fragmented landscapes. Con-servation Biology 19:768–782.

He, J-S., Q-B. Zhang, and F. A. Bazzaz. 2005. Differ-ential drought responses between saplings andadult trees in four co-occurring species of NewEngland. Trees 19:442–450.

Hobbie, E. A. 2005. Using isotopic tracers to followcarbon and nitrogen cycling of fungi. In J.Dighton, P. Oudemans, and J. White (eds.), TheFungal Community: Its Organization and Role in theEcosystem. CRC Press, Boca Raton, Florida.

———. 2006. Carbon allocation to ectomycorrhizalfungi correlates with total belowground alloca-tion in culture studies. Ecology 87 3:563–569.

Hobbie, E. A. and H. Wallander. 2006. Integrating

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ectomycorrhizal fungi into quantitative frame-works of forest carbon and nitrogen cycling. InG. M. Gadd (eds.), Fungi in Biogeochemical Cycles.Cambridge University Press, Cambridge.

Hobbie, J. E. and E. A. Hobbie. 2006. DecodingNature’s signals: 15N content in symbiotic fungiand plants estimates nitrogen and carbon fluxrates in an Arctic ecosystem. Ecology 87, 4,816–822.

Horii, C. V., S. C. Munger, S. C. Wofsy, M. Zahnis-er, D. Nelson, and J. B. McManus. 2004. Fluxesof nitrogen oxides over a temperate deciduousforest. Journal of Geophys. Res. 109 D08305,doi:10.1029/2003JD004326.

———. 2005. Atmospheric reactive nitrogen concentra-tions and flux budgets at a northeastern U. S. for-est site. Agricultural and Forest Meteorology133:210–225.

Hou, J., Y. Huang, Y. Wang, B. Shuman, W. W.Oswald, E. Faison, and D. R. Foster. 2006. Postglacial climate reconstruction based on compound-specific D/H ratios of fatty acids fromBlood Pond, New England. Geochemistry, Geo-physics, Geosystems Q03008.

Ingerson, A. E. 2004. From contracts to conse-quences: History as an adaptive managementtool for national forests and local communities.Pp. 65–85 in Forest Conservation and Stewardship in Massachusetts. C. H. W. Foster and P. R. Hagenstein (Eds.). Harvard Forest, Petersham,MA.

Jefts, S. and D. Orwig. 2005. The effects of HWAoutbreaks on ecosystem level changes in south-ern New England. Pp. 305–306 in Proceedings ofthe third symposium on hemlock woolly adelgid in the eastern United States. Reardon, R. and B. Onken(Eds.). USDA Forest Service, Newtown, PA.

Jones, J. A. 2006. Intersite comparisons of rainfall-runoff processes. In pp 1839–1854 M. Anderson(Ed.), Encyclopedia of Hydrologic Sciences. John Wiley,Chichester, UK.

Kelty, M. J. and A. W. D’Amato. 2006. Historical per-spective on diameter-limit cutting in northeasternforests. In: Diameter-limit Cutting in NortheasternForests 2005 May 23–24. Gen. Tech. Rep. NE-341, Newtown Square, PA.

Kittredge, D. B. 2004. Defining a Vision for Massa-chusetts Forest. Pp. 19–24 in Forest Conservationand Stewardship in Massachusetts. C. H. W. Fosterand P. R. Hagenstein (Eds.). U.S.A.

———. 2004. Extension/outreach implications for

America’s family forests owners. Journal of Forestry102:15–18.

———. 2004. Forests and forestry of Massachusetts. Pp.11–18 in Forest Conservation and Stewardship in Mass-achusetts. C. H. W. Foster and P. R. Hagenstein(Eds.). U.S.A.

———. 2005. The cooperation of private forest ownerson scales larger than one individual property:international examples and potential applicationin the United States. Forest Policy and Economics7:671–688.

———. 2005. The cooperation of private forest ownerson scales larger than one individual property:international examples and potential applicationin the United States. Forest Policy and Economics7:671–688.

Kittredge, D. B., K. Clark, M. Ohmann, P. Huckery,and T. French. 2006. Protection of habitat forstate-listed rare flora and fauna in Massachusettsduring timber harvesting. Natural Areas Journal26:198–207.

Köster, D. and R. Pienitz. 2006. Seasonal DiatomVariability and Paleolimnological Inferences — ACase Study. Journal of Paleolimnology 35:395–416.

Köster, D., R. Pienitz, B. B. Wolfe, S. Barry, D. R.Foster, and S. S. Dixit. 2005. Paleolimnologicalassessment of human-induced impacts on thenutrient balance of Walden Pond (Massachusetts,U.S.A.) during the last three centuries usingdiatoms and stable isotopes. Aquatic EcosystemHealth and Management 8:117–131.

Lai, C. T., J. R. Ehleringer, A. J. Schauer, P. P. Tans,D. Y. Hollinger, K. T. Paw U, J. W. Munger, andS. C. Wofsy. 2005. Canopy-scale delta 1(3)C ofphotosynthetic and respiratory CO2 fluxes:observations in forest biomes across the UnitedStates. Global Change Biology 11:633–643.

Levitt, J. N. (Editor) 2005. From Walden to Wall Street:Frontiers of Conservation Finance. Island Press andLincoln Institute.

Lindenmayer, D. B., D. R. Foster, J. F. Franklin , M.L. Hunter, R. F. Noss, F. A. Schmiegelow, and D.Perry. 2004. Salvage harvesting policies after nat-ural disturbance. Science 303:1303.

Magill, A. H., J. D. Aber, W. S. Currie, K. J. Nadel-hoffer, M. E. Martin, W. H. McDowell, J. M.Melillo, and P. Steudler. 2004. Ecosystemresponse to fifteen years of chronic nitrogen addi-tions at the Harvard Forest LTER, Massachu-setts, U.S.A. Forest Ecology and Management196:7–28.

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Mansfield, C., S. Pattanayak, W. McDow, R. I.McDonald, and P. N. Halpin. 2006. Shades ofgreen: Measuring the value of urban forests inthe housing market. Journal of Forest Economics11(3):177–199.

McDonald, R. I. 2006. A world of the city, by thecity, for the city. In J. Harf, M. Lombardi (eds.).Taking Sides: Clashing views in global issues. McGraw-Hill, New York.

McDonald, R. I. and D. L. Urban. 2006. Spatiallyvarying rules of landscape change: Lessons froma case study. Landscape and Urban Planning74:7–20.

———. 2006. Forest edges and forest composition inthe North Carolina Piedmont. Biological Invasions8:1049–1060.

McDonald, R. I., G. Motzkin, M. S. Bank, D. B. Kit-tredge, J. Burk, and D. R. Foster. 2006. Forestharvesting and deforestation relationships overtwo decades in Massachusetts. Forest & EcologyManagement 227:31–41.

McDonald, R. I., M. McKnight, D. Weiss, E. Selig,M. O’Connor, C. Violin, and A. Moody. 2005.Species compositional similarity and ecoregions:Do ecoregion boundaries represent zones of highspecies turnover? Biological Conservation 126:24–40.

McDowell, W. H., A. Magill, J. A. Aikenhead-Peter-son, J. D. Aber, J. L. Merriam, and S. S. Kaushal.2004. Effects of chronic nitrogen amendment ondissolved organic matter and inorganic nitrogensoil solution. Forest Ecology and Management196:29–41.

McLachlan, J. S., J. S. Clark, and P. S. Manos. 2005.Molecular indicators of tree migration capacityunder rapid climate change. Ecology 86:2088–2098.

Micks, P., J. D. Aber, R. D. Boone, and E. A. David-son. 2004. Short-term soil respiration and nitro-gen immobilization response to nitrogen applica-tions in control and nitrogen-enriched temperateforests. Forest Ecology and Management 196:57–70.

Micks, P., M. R. Downs, A. H. Magill, K. J. Nadel-hoffer, and J. D. Aber. 2004. Decomposing litteras a sink for 15N-enriched additions to an oak for-est and a red pine plantation. Forest Ecology andManagement 196:71–87.

Moir, R. 2004. Shared land planning and manage-ment in Massachusetts with the National ParkServices, U.S. Fish and Wildlife Service, and pub-lic-private partnerships. Pp. 33–50 in Forest Con-

servation and Stewardship in Massachusetts. C. H. W.Foster and P. R. Hagenstein (Eds.). Harvard For-est, Petersham, MA.

Moore, G. W., B. J. Bond, J. A. Jones, N. Phillips, andF. C. Meinzer. 2004. Structural and composi-tional controls on transpiration in a 40- and 450-yr-old riparian forest in western Oregon, USA.Tree Physiology 24:481–491.

Munger, J. W., C. Barford, and S. C. Wofsy. 2004.Exchanges between the Forest and Atmosphere.Pp. 202–233 in Forest Landscape Dynamics in NewEngland, Ecosystem Structure and Function as a Conse-quence of History, D. Foster and J. Aber, (Eds.). YaleUniversity Press, New Haven.

Nadelhoffer, K. J., B. P. Colman, W. S. Currie, A. H.Magill, and J. D. Aber. 2004. Decadal scale fatesof 15N tracers added to oak and pine stands underambient and elevated N inputs at the HarvardForest (U.S.A.). Forest Ecology and Management196:89–107.

Ollinger, S. V. 2005. Biodiversity and EcosystemFunction in Forests. Review of: Scherer-Loren-zen, Michael, Christian Körner, and Ernst-DetlefSchulze, editors. 2005. Forest diversity and function:temperate and boreal systems. Springer, New York.Ecology 86:2256–2257.

Ollinger, S. V. and M. L. Smith. 2005. Net primaryproduction and canopy nitrogen in a temperateforest landscape: an analysis using imaging spec-trometry, modeling and field data. Ecosystems8:760–778.

Orwig, D. A. and D. B. Kittredge. 2005. Silviculturaloptions for managing hemlock forests threatenedby hemlock woolly adelgid. Pp. 212–217 in Pro-ceedings of the third symposium on hemlock woolly adelgidin the eastern United States. Reardon, R. and B.Onken (Eds). USDA Forest Service, Newtown,PA.

Osterweil, L. J., A. Wise, L. Clarke, A. M. Ellison, J.Hadley, E. R. Boose, and D. R. Foster. 2006.Process technology to facilitate the conduct of science. In: International Software Process Workshop.Springer-Verlag, Germany.

Primack, R. and R. Corlett. 2005. Tropical Rainforests:An Ecological and Biogeographical Comparison. Black-well Science.

Putz, F. 2004. Hope and the hemlock. Sanctuary43:10–11.

Ratchford, J. S., S. E. Wittman, E. S. Jules, A. M. Elli-son, N. J. Gotelli, and N. J. Sanders. 2005. Theeffects of fire, local environment, and time on ant

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assemblages in fens and forests. Diversity and Dis-tributions 11:487–497.

Richardson, A. D., A. S. Bailey, E. G. Denny, C. W.Martin, and J. O’Keefe. 2006. Phenology of anorthern hardwood forest canopy. Global ChangeBiology 12:1174–118.

Ross, K. 2004. Using forest conservation easementsproactively to encourage stewardship. Pp. 25–32in Forest Conservation and Stewardship in Massachu-setts. C. H. W. Foster and P. R. Hagenstein (Eds.).Harvard Forest, Petersham, MA.

Sack, L., C. M. Streeter*, and N. M. Holbrook. 2004.Hydraulic analysis of water flow through leavesof sugar maple and red oak. Plant Physiology 134:1824.

Sevanto, S., N. M. Holbrook, M. Zwieniecki, T. N.Mikkelsen, W. J. Munger, and S. Wofsy. 2005.Diurnal pattern in water tension of tree stemsbegins before budbreak and does not cease withleaf fall. In: The 90th Annual Meeting of The Ecolog-ical Society of America (ESA) and IX International Con-gress of Ecology (INTECOL).

Sevanto, S., T. Hölttä, A. Riikonen, E. Nikinmaa, T.N. Mikkelsen, W. J. Munger, S. C. Wofsy, T.Vesala, and N. M. Holbrook. Information onsugar and water circulation patterns of trees bydiurnal diameter variation detection. In: ReportSeries in Aerosol Science, 1st iLEAPS Science Conference,21–26 Jan 2006.

Sims, D. A., A. F. Rahman, V .D. Cordova, D. D.Baldocchi, L. B. Flanagan, A. H. Goldstein, D. Y.Hollinger, L. Misson, R. K. Monson, H. P.Schmid, S. C. Wofsy, and L. K. Xu. 2005. Mid-day values of gross CO2 flux and light use effi-ciency during satellite overpasses can be used todirectly estimate eight-day mean flux. Agriculturaland Forest Meteorology 131:1–12.

Sodhi, N. S., L. H. Liow, and F. A. Bazzaz. 2004.Avian extinctions from tropical and subtropicalforests. Annual Review of Ecology, Evolution and Systematics 35: 323–345.

Spicer, R. and N. M. Holbrook. 2005. Within-stemoxygen concentration and sap flow in four tem-perate tree species: does long-lived xylemparenchyma experience hypoxia? Plant, Cell andEnvironment 28, 192–201.

Stadler, B., T. Muller, and D. Orwig. 2006. The ecol-ogy of energy and nutrient fluxes in hemlockforests invaded by the hemlock woolly adelgid.Ecology 87:1792–1804.

Stadler, B. T. Muller, D. Orwig, and R. Cobb. 2004.

Hemlock woolly adelgid: canopy impacts trans-forming ecosystem processes and landscapes.Ecosystems 8: 233–247.

———. 2005. Hemlock Wooly Adelgid in New Eng-land Forests: Canopy impacts transformingecosystem processes and landscapes. Ecosystems8:233–247.

Stinson, K.A. 2004. Natural selection favors rapidreproductive phenology in Potentilla pulcherrima(Rosaceae) at opposite ends of a subalpinesnowmelt gradient. American Journal of Botany91:531–539.

Stinson, K. A. 2005. Effects of snowmelt timing andneighbor density on the distribution of the highaltitude plant, Potentilla diversifolia. Arctic, Antarcticand Alpine Research 37:379–386.

Stinson, K. A. and F. A. Bazzaz. 2006. CO2-enrich-ment reduces reproductive dominance in com-peting stands of Ambrosia artemisiifolia (commonragweed). Oecologia 146:155–163.

Stinson, K. A., J. Tran*, J. Petzold*, and F. A. Bazzaz.2006. Photosynthetic and architectural responsesof common ragweed to elevated CO2 result indecreased asymmetry of light competition. GlobalChange Biology 12.

Stinson, K. A., S. A. Campbell, J. R. Powell, B. E.Wolfe, R. M. Callaway, G. C. Thelen, S. G. Hal-lett, D. Prati, and J. N. Klironomos. 2006. Inva-sive plant suppresses the growth of native treeseedlings by disrupting belowground mutu-alisms. Public Library of Science, Biology 4:e140.

Sudderth, E. A., K. A. Stinson, and F. A. Bazzaz.2006. Plant-mediated aphid population responseto elevated CO2 and increased nitrogen availabil-ity. Global Change Biology 11:1–12.

Sulzman, E., J. Brant, R. D. Bowden, and Lajtha, K.2005. Contribution of aboveground litter, below-ground litter, and rhizosphere respiration to totalsoil CO2 efflux in an old growth coniferous. Bio-geochemistry 73:231–256.

Taverna, K., D. L. Urban, and R. I. McDonald.2005. Modeling landscape vegetation pattern inresponse to historic land use: a hypothesis-drivenapproach for the North Carolina Piedmont,USA. Landscape Ecology 46.

Tomlinson, P. B. 2006. Stem anatomy of climbingpalms in relation to long distance water trans-port. Aliso 22:263–275.

———. 2006. The uniqueness of palms. Botanical Jour-nal of the Linnean Society 151:5–14.

Tomlinson P. B., J. B. Fisher, F. Hallé, and R. Vil-

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lalobos. 2005. Development of woody branchattachments in Schefflera (Araliaceae or Apiaceae).American Journal of Botany 92:1765–1773.

Tomlinson, P. B. and J. B. Fisher. 2005. Developmentof non-lignified fibers in leaves of Gentumgnemon (Gnetales). American Journal of Botany92:383–389.

Urban, D. L., R. I. McDonald, E. S. Minor, and E.A. Treml. 2006. Causes and consequences ofland use change in the North Carolina Piedmont.Pp. 239–258 In J. Wu, B. Jones, H. Li, and O. L.Loucks (eds.), Scaling and Uncertainty Analysis inEcological Studies. Springer, Dordrecht, TheNetherlands.

Venterea, R. T., P. M. Groffman, L. V. Verchot, A. H.Magill, and J. D. Aber. 2004. Gross nitrogenprocess rates in temperate forest soils exhibitingsymptoms of nitrogen saturation. Forest Ecologyand Management 196:129–142.

Von Holle, B. 2005. Biotic resistance to invaderestablishment of a southern Appalachian plantcommunity is determined by environmental con-ditions. Journal of Ecology 93:16–26.

Von Holle, B. and D. Simberloff. 2004. Testing Fox’sassembly rule: Does plant invasion depend uponrecipient community structure? Oikos 105:551–563.

Von Holle, B., K. A. Joseph*, R. G. Lohnes*, and E.F. Largay*. 2006. Facilitations between the intro-duced nitrogen-fixing tree, Robinia pseudoacacia,

and nonnative plant species in the glacial out-wash upland ecosystem of Cape Cod, Massachu-setts. Biodiversity and Conservation 15:2197–2215.

Wakefield, A. E., N. J. Gotelli, S. E. Wittman, and A.M. Ellison. 2005. The Effect of Prey Additionalters nutrient stoichiometry of the carnivorousplant Sarracenia Purpurea. Ecology 86:1737–1743.

Wender*, N. J., C. Polisetty, and K. Donohue. 2005.Density-dependent processes influencing the evo-lutionary dynamics of dispersal: A functionalanalysis of seed dispersal in Arabidopsis thaliana(Brassicaceae). American Journal of Botany 92:960–971.

Whigham, D. F. 2004. Ecology of woodland herbs intemperate deciduous forests. Annual Review ofEcology, Evolution, and Systematics 35:538–621.

Xiao, X., Q. Zhang, S. Saleska, L. Hutyra, P. DeCamargo, S. Wofsy, S. Frolking, S. Boles, M.Keller, and B. Moore III. 2005. Satellite-basedmodeling of gross primary production in a sea-sonally moist tropical evergreen forest. RemoteSensing of the Environment 94:105–122.

Yi, C., R. Li, P. S. Bakwin, A. Desai, D. M. Ricciuto,S. P. Burns, A. A. Turnipseed, S. C. Wofsy, J. W.Munger, K. Wilson, and R. K. Monson. 2004. Anonparametric method for separating photosyn-thesis and respiration components in CO2 fluxmeasurements, Geophysical Research Letters 31, No.17.

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Simes Tract.

ACKNOWLEDGMENT OF SUPPORT

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

AnonymousFriends of the Harvard Forest Andrew W. Mellon FoundationCardinal Brook TrustCommonwealth of Massachusetts

Dept of Environmental ManagementDivision of Fisheries and Wildlife Natural Heritage & EndangeredSpecies Program

Department of Energy, National Institutefor Global Environmental Change

Conservation and Research FoundationDoris Duke Charitable FoundationGeorge & Jane Mifflin Memorial FundGreenleaf FoundationHarvard University

Center on the EnvironmentDavid Rockefeller CenterThe Clark FundThe Cooke Fund

Highstead Foundation, Inc.Massachusetts Environmental TrustNational Park FoundationNational Park ServiceNational Science Foundation

Digital Government ProgramEcosystem StudiesEd En Venture FundInternational ProgramsInformation Technology Research Long-term Ecological StudiesEcology ProgramResearch Experience for Undergraduates

Sweetwater TrustThe Cabot Family Charitable TrustThe Fields FoundationThe Fine Family TrustThe Guilford FundThe Nature ConservancyUSDA Forest Service

GIFTS

The Harvard Forest library and archives thanks thefollowing for gifts received this past year: James Bairddonated the following books: Vegetation of New Jersey,The Flora of the Connecticut Arboretum, A Textbook of Sys-tematic Botany, The Vegetation of Wisconsin, Manual of Veg-etation Analysis and 2004 journals issues of The Auk,Ornithological Monographs, and The Journal of FieldOrnithology. A special thanks also to Jim for also pass-ing along copies of rare early Harvard Forest bul-letins. Mrs. Solon Barraclough donated researchnotes from her husband’s work at the Harvard For-est during the 1940s and 1950s. Jiquan Chen, a for-mer Bullard Fellow, donated a copy of his recentbook Nature’s Giving — Ecology of Ecosystems and Land-scapes. Charles W. H. Foster donated four books he

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wrote in the Experiments in Bioregionalism series:The New England River Basins Story, The Cape CodNational Seashore, The Appalachian National Scenic Trail,and Yankee Salmon. Mr. Francis Holmes donated alarge collection of books and reprints on old floras ofMassachusetts, tree disease guides, and storm distur-bance reports. David Tatlock donated a copy of amap of the former Fisher family property on ProspectHill and more photographs and letters from his fam-ily to add to our R. T. Fisher archival files. A dissect-ing microscope and binocular stage microscope withmultiple optics and accessories, along with a series ofbotanical books were donated by Mary Ann Walker.Historical researcher Electa Tritsch donated copies ofher report on the history of the Trustees of Reserva-tions properties in Petersham.

NEW FUNDING 2005

David Foster and David Orwig — Nutrient AnalysisEquipment for Community, Ecosystem, Hydrologi-cal and Physiological Research at Harvard Forest;NSF REU Supplement DBI-0521263: $10,005.

David Foster and Julian Hadley — Effects of ForestAge, Soil Drainage and Interannual Climate Varia-tion on Deciduous Forest Carbon Exchange: A Com-parison to Long-term Harvard Forest Measurements,in a Contrasting Forest Type; DOE NIGEC 011213-HU: $362,680.

Aaron Ellison and Kathleen Donohue — REU Site:Harvard Forest REU Program in Forest Ecology2005–2009: Multi-Scale Investigations of a ForestedEcosystem in a Changing World; NSF 0452254:$735,042.

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Aaron Ellison — Collaborative Research: Effects ofNutrient Stress on a Co-evolved Food Web; NSFDEB-05-28625, RET Supplement for NSF 02-35128: $10,560.

Aaron Ellison — Collaborative Research: Effects ofNutrient Stress on a Co-evolved Food Web; NSFDEB-05-20792, REU Supplement for NSF 02-35128: $7,000.

Aaron Ellison — FSML: Infrastructure for Whole-Plant Biology and Experimental Plant Ecology at theHarvard Forest; NSF DBI-05-20794, REU Supple-ment for NSF 03-30605: $10,005.

David Foster and David Orwig — Landscape Patternsof Hemlock Wooly Adelgid Damage in New EnglandForests NSF REU Supplement DEB-0236897:$6,000.

NEW FUNDING 2006

Aaron Ellison — Moths, ants, and carnivorous plants:the spatial dimension of species interactions. NSFDEB 05-41680. $585,000.

Aaron Ellison — FSML: Infrastructure for Whole-Plant Biology and Experimental Plant Ecology at theHarvard Forest. NSF DBI-05-20794, REU Supple-ment for NSF 03-30605: $10,691.

David R. Foster — Proposal for a Workshop to MakeNEON – A Transformational Ecological ResearchProgram that Robustly Embraces the HumanDimension held in fall 2005 at Harvard Forest. DBI-0525165. $28,703.

David Foster, Betsy Colburn, and Pamela Snow —Harvard Forest Freshwater Ecology Research in theSchoolyard, MET 5094913-01 $15,200.

David Foster and David Orwig — Nutrient AnalysisEquipment for Community, Ecosystem, Hydrologi-cal, and Physiological Research at Harvard Forest;NSF REU Supplement DBI-0521263: $6,000.

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David R. Foster Petersham, MassachusettsDirector December 2006

New maintenance garage on Prospect Hill.