Land-Use History (1730-1990) and Vegetation Dynamics in Central New England,... Author(s): David R. Foster
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Land-Use History (1730-1990) and Vegetation Dynamics in Central New England,... Author(s): David R. Foster
Land-Use History (1730-1990) and Vegetation Dynamics in Central New England, USA Author(s): David R. Foster Source: Journal of Ecology, Vol. 80, No. 4 (Dec., 1992), pp. 753-771 Published by: British Ecological Society Stable URL: http://www.jstor.org/stable/2260864 . Accessed: 03/09/2013 16:19 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. . British Ecological Society is collaborating with JSTOR to digitize, preserve and extend access to Journal of Ecology. http://www.jstor.org This content downloaded from 128.103.149.52 on Tue, 3 Sep 2013 16:19:33 PM All use subject to JSTOR Terms and Conditions Journal of Ecology 1992, 80, 753-772 Land-use history(1730-1990) and vegetation dynamicsin centralNew England,USA DAVID R. FOSTER Harvard Forest,Harvard University, Petersham,MA 01366, USA Summary 1. Historiesofchangingland use and vegetationof a 380-haforestedarea in central Massachusetts(Prospect Hill tractof the Harvard Forest) were reconstructedto investigate(i) theenvironmental controlsoverland ownershippatterns,agricultural practice and loggingactivity,and (ii) the vegetationresponse to these land-use factors. 2. Forest clearance and agriculturalexpansion parallel trendsfor central New England: increasingratesof deforestationthroughthe late eighteenthcenturyled to a peak in 1820-80 when more than 80% of the land was open. Reforestation on abandoned fieldscommencedin 1850 and increasedprogressively throughthe earlytwentiethcentury. 3. Ownershippatternsvaried temporallyin turnoverrate and size of individual holdings.Twenty-five lots comprisingthe studyarea were sold an average of 13 timesin theperiod1730-1910. Land sales weregreatestin theperiodofspeculation and low-intensity agriculture(1730-90), lowestduringthetransition to commercial agricultureand small-scaleindustry(1790-1840), and high duringthe period of agriculturaldecline in the mid to late 1800s. 4. Land use in the mid 1800s, includingwoodlot (13% of the studyarea), tilled fields(16%), pasture (70%) and marsh (1%), formedan intricatepatternbest explained by soil drainage and proximityto farmhousesand town roads. This land-use pattern controlled the reforestationprocess: field abandonment and reforestationproceeded outward from poorly drained pasture adjacent to the continuouswoodlotsand eventuallyincludedproductivetilledland. 5. The consequences of 250 years of land-use activityvary at differentscales. Regionally,the distribution of modernand pre-settlement foresttypesmatchwell despitestructuralchangesand the loss of some tree species. At a landscape scale, are stronglycontrolledby land use. Canonical corremodernforestcharacteristics variationis bestexplainedbyhistorical spondenceanalysisindicatesthatcommunity factors(distinctionbetweenprimaryand secondarywoodlands,forestage, cutting historyand timingof site abandonment)and site factors(slope positionand soil drainage). Picea rubens and Tsuga canadensis forestsare restrictedto primary woodlands, Pinus strobusand sproutsof Castanea dentataare largelyconfinedto old pastures, and Betula populifolia, Populus spp. and Acer rubrumare most abundantin cut-overold-fieldPinus stands. 6. Long-termforesttrendsin the twentiethcenturyinclude a decrease in the importanceof Pinus strobusdue to loggingand the 1938 hurricane,a gradual decline in early successionalhardwoods (Betula populifolia,Populus spp., Acer rubrum),and increase in later successionalspecies (Quercus rubra, Q. velutina, Acer saccharum). Tsuga and northern-hardwood species (Acer saccharum,Fagus grandifolia)declineddramaticallythroughoutthe settlementperiod; however,the major foresttrendover the past 100 yearshas been a continualincreasein Tsuga. Key-words:agriculturalhistory,canonicalcorrespondenceanalysis,deforestation, forestdynamics,humandisturbance 753 Journalof Ecology 1992,80, 753-772 This content downloaded from 128.103.149.52 on Tue, 3 Sep 2013 16:19:33 PM All use subject to JSTOR Terms and Conditions 754 Land use and vegetationhistory in New England Introduction Despite the long traditionin north-western Europe of utilizingpalaeoecological and historicalmethods to investigatehuman impactson the environment and vegetation(Iversen 1973; Peterken & Game 1981; Peglar, Fritz & Birks 1988), there are few comparable studies in NorthAmerica. This difference is oftenattributedto the shorterdurationof intensivehumanimpactin much of NorthAmerica versusEurope (c. 400 yearsvs. >5000 years) but is nonethelessremarkablegiventhe intensity and geographicalextentof humanimpactin theNew World and the ecological importanceof understanding this history(Marsh 1864; Fisher1933; Ecological Society of America 1991). Investigations of land-usehistory and vegetationchange provide: 1. a backgroundforunderstanding thedevelopment of the modern vegetational landscape, which is essentialforanyecologicalstudy(Christensen1989); 2. informationon the response of communities to novel disturbanceprocesses or intensitiesthat may be comparedwithstudiesof naturalprocesses to evaluate the resistance or resilience of communities to contrastingfactors (Houghton et al. 1983; Schoonmaker& Foster 1991); 3. perspectives thatmaybe usefulin thepreservation of unique communitiesor managementof cultural landscapes (Birks et al. 1988). Conclusive studies linking human activityand ecosystemchange require comprehensiveanalyses ofland-usehistoryand long-term vegetationrecords. The presentstudyseeks to evaluate thesetwo kinds of information for a nearly400-ha forestedarea in central Massachusetts(Prospect Hill tract of the Harvard Forest) and for a period extendingfrom the mid eighteenthcenturyto the present. The studyhas three major objectives: (i) to document spatial and temporal patternsof land-use and to relate these to environmentalfactorsand cultural conditions;(ii) to identifythe major environmental and humanfactorsthatare responsibleforchanges in the structureand compositionin the vegetation duringthisperiod, and (iii) to documentlong-term changesof the vegetationin thisstudyarea. Although the forest investigatedis unusual in having an extensively documented history, the major patternsof land-use and change in forest cover are representativeof broad upland areas of the north-eastern USA (Bidwell 1916; McKinnon, Hyde & Cline 1935; Black & Brinser1952). In order to place the resultsof the research in a broader geographicalframework,the study begins with a descriptionof the early post-settlement vegetation ofthetownshipofPetershamand WorcesterCounty, Massachusetts.It thenexaminesthe specifichistory and effectsof land-usepracticeson theProspectHill tractof the HarvardForest in Petersham. Studyarea PHYSICAL SETTING Worcester County occupies 3900km2 in central Massachusetts(Fig. 1). The westerntwo-thirdsof the county formsan undulatingupland generally exceeding 250 m a.s.l., whereas to the east and south-eastthe land slopes to a prevailingaltitude of 100m. Second-growthforest currentlycovers approximately70% of the land area (MacConnell & Niedzweidz 1974). Petershamis located in north-western Worcester Countyat an average altitudeof 275 m a.s.l. Relief of c. lOOm encompasses a series of north-south trendingridges and valleys. The soils are largely acidic and derivedfromgranodioritesand gneisses. The meanannualtemperature is 8 5 ?C, thefrost-free season averages 5 monthsand the annual precipitation is 105cm including150cm of snow (Rasche 1953). The townshipis 90% forested. The Prospect Hill tract of the Harvard Forest comprises380ha of the northern, highestportionof the major ridgein Petersham.Altituderangesfrom 270 to 420m a.s.l., steep slopes occur towardsthe westernand north-eastern marginsof the tract,and mostof thearea is undulating(Fig. 2). Variabilityin I 72 W I I 680 46"N- Northern Hardwoods . and Spruce- Fir TransitionHardwoods ""HARVARD FOREST 420 Pitch Pine-Oak / CentralHardwoods km Fig. 1. Forest vegetationmap of New England showing WorcesterCounty, Massachusetts(WC) extendingfrom the Central Hardwood zone throughto the Northern Hardwoodzone. The locationof theHarvardForestin Petersham,Massachusettsin indicatedby the solid circle. ModifiedfromWestveld(1956). This content downloaded from 128.103.149.52 on Tue, 3 Sep 2013 16:19:33 PM All use subject to JSTOR Terms and Conditions 755 D.R. Foster (a) (b PrspectHill 40 SA/I~~~~~ Sanderson Farmhouse U TanyardMillSite (c) i---'-~~~~~~~~~~~ \/' ,rir , Fig. 2. The Prospect Hill area of the Harvard Forest showing (a) topography (5-m contours), (b) soil drainage (from very-well drained (light shading) to very-poorly drained (black) and (c) location of stonewalls and roads. The maps are at matchingscales and the outline of the soils map conformsto that of the studyarea boundary. Locations of the John Sanderson farmhouse, tanyard mill site and top of Prospect Hill are indicated on all maps.\ \ ,-_ L L_ ~ ' ' ~ '~ '__ , , ',X1 ,_, ,, '', 8 " .'\ ,' - --- l , s 5 , '----- Stonewalls \~Roads relief,depthto bedrock,and presenceof a fragipan create a highlydissected patternof soil drainage population density (Figs 3 and 4; Torbert 1935; Pabst 1941; Barraclough1949). This historycan be divided intofivemajor periods: 2b). (Fig. 1730-50 speculation; 1750-90 low-intensity agriculture; CULTURAL SETTING 1790-1850 commercialagriculture and smallindustry; Duringthe past 250 yearsthe centralMassachusetts 1850-1920 farmabandonmentand industrialization; landscape has undergone several transformations 1920-90 residentialperiod in response to changes in land-use practices and (Raup & Carlson 1941; Black & Wescott1959). This content downloaded from 128.103.149.52 on Tue, 3 Sep 2013 16:19:33 PM All use subject to JSTOR Terms and Conditions 756 Land use and vegetationhistory in New England 100 50 ~0 0 . 1720 1760 1800 1840 1880 1920 1960 Year Fig. 3. Historicalchanges in forestcover forthe State of Massachusetts(o), the townshipof Petersham(0), and the ProspectHill tract(U). Sources of information include Dickson & McAfee (1988), MacConnell (1975), Rane (1908) and Baldwin (1942) for Massachusetts,and Raup & Carlson (1941), Anonymous (1959), MacConnell & Niedzwiedz (1974), Averill, Averill & Stevens (1923), Cook (1917) and Rane (1908) forPetersham.Estimatesof forestcover on the ProspectHill tractwere derivedfrom land-use notes in deeds, Spurr (1950) and unpublished sourcesin the HarvardForest Archives. F s rv ( -)dro r l The townshipof Petershamwas settledin 1733. With limited access to markets, the eighteenthcenturyrural economy was based on farmingand small-scalecommercialactivity(Coolige 1948; Pruitt 1981; Baker & Patterson 1986). Forest clearance proceeded initiallyat a pace of 1-4% per year and increasedde(Fig. 3). Improvedtransportation mandforagricultural productsin the late eighteenth and earlynineteenthcenturiespromptedan increase in commercialfarmingand small-scaleindustryin rural centralNew England (Pabst 1941; Thorbahn & Mrozowski 1979; Rothenberg 1981). Forest clearance to provide pasture land for beef cattle and sheep resulted in an increase in open land fromapproximately50% of the townshipin 1800 to nearly85% in 1850 (Figs 3 and 4). Remaining forestsoccupied steep and rockyslopes, wetlandsor narrowvalleys and were cut for timber,fuelwood and tanbark and were often grazed (Cline et al. 1938; Gould 1942; Spurr1950). 19985; Fig. 4. The townshipof Petershamdepictingforestedareas (black) and open land (white) in 1830, 1900, 1938 and 1985. In the map for 1900 the stipplingindicatesformeragriculturalland abandoned fromc. 1870 to c. 1900 thatseeded into old-fieldPinus strobus.Progressiveabandonmentof farmsled to a concentrationof non-forestedlands on north-south ridges.Data were obtainedfromtheAtlas of WorcesterCounty(1830, unpublished),unpublishedmaps fromthe Harvard Forest Archives(1935-39) and air-photograph analysis(1985; 1:24,000colour-infra-red). This content downloaded from 128.103.149.52 on Tue, 3 Sep 2013 16:19:33 PM All use subject to JSTOR Terms and Conditions 757 D.R. Foster V (b) Fig. 5. (a) Millstone,streamand retaining wall of an abandoned mill on the Prospect Hill tractoftheHarvardForestin Petersham, Massachusetts. The mill was activelyused from 1780 to 1850 to grindTsuga barkhs and fomrexetoffedsetndtrug forests~~ doiae ~ to~produce yQerutannic ~ acid ur,Ae Castanea wood foruse in leathertanning.Providinga major source of income supplementalto agriculturalac, the mill exerteda pronouncedimpact tivity on thewoodlandsin theneighbouring region rubrum as they were Betu)Mlasoe cutpaprieram to provide Freaxinungs a frequentlyand of tannin. wamerfcana asandhere atlo The themodern Prospect Hillforest trct regularsource is comprisedofForest.Poorpsb Acer saccharumand HltrcofteHarvard M.Fraxinus Fluetsha) americana established in the late 1800s. (b) Broad-scale abandonmentof agricultural land across New England in the mid to late 1800sgave rise to extensiveareas of secondgrowth forest. Stone walls delimitingthe This content downloaded from 128.103.149.52 on Tue, 3 Sep 2013 16:19:33 PM All use subject to JSTOR Terms and Conditions 758 Land use and vegetationhistory in New England Commencingin the mid 1800s the populationof Petershamdeclined as youngerinhabitantsleftfor urbanjobs or agriculturalopportunitiesin the Midwest (Fisher 1921; Black & Brinser 1952). Farms wereabandonedand neglectedfieldswerereforested (Fig. 5) (Munyon1978; Garrison1987). The modern period has includeda nearlycompletecessation of agriculturalactivity,broad-scaleregrowthof forest cover and a conversionof the townto a residential status.Since the 1930stherehas been an increasein homes and a reversalin the downwardpopulation of directhuman trendbut a decrease in theintensity impacton the land. Methods VEGETATION OF WORCESTER COUNTY Data on the early post-settlement(c. 1770-90) vegetationof WorcesterCountywere derivedfrom Peter Whitney's (1793) account which includes a geographical descriptionof each township,its landscape and vegetation. Whitneytravelled the within20-30 yearsof the date of countyextensively incorporation ofmostofthetownshipsand described the vegetationbefore extensivehuman alteration. Whitney's data were analysed using detrended correspondenceanalysis(Hill & Gauch 1980) with species abundance assigned as either present or abundant (see Table 1). Results of the samples (townships)ordinationwere used to identifyforest type groupingsand to develop a countyvegetation map. A phytosociologicaltable was constructed fromsamples and species output.The map of early forestvegetationwas compared to recentregional forest descriptions(Westveld 1956). Species' nomenclaturefollowsFernald (1973). The long-termimpact of land-use on presentday forestcompositionwas analysed in two ways. Using a GeographicInformationSystemthe spatial overlap between soil drainage, land-use history and foresttypes in 1908 was examined and transitions in forestcommunitiesbetween 1908, 1937 and 1986 were documented. Informationon soils and vegetationwas derived fromHarvard Forest (HF) Archives.Canonical correspondenceanalysis (CANOCO, terBraak 1986) was thenused to evaluate the relationshipbetween the vegetation and environmentalvariables (includingsite and historical factors)forthe forestin 1937. The 1937 vegetation was selected for intensiveanalysis because of the availabilityof an extensive data set and the fact that the 1937 vegetation representedthe maximum extentof secondarydevelopmentbefore the 1938 hurricane,when approximately75% of the timber was windthrown(Foster 1988; Foster & Boose 1992). Vegetational informationanalysed in CANOCO, included 253 samples (approximately 0-04-haplots) and 105 species. Tree species abunscale based on dance was assigned on a five-factor volume, whereas shrubs, herbs and mosses were recordedas presentor absent.Sitevariablesanalysed includedsoil drainage(fiveclasses; Spurr1950) and slope position(fiveclasses). Historicaland land-use variablesincludedstandage (derivedfromthe 1937 survey), time since agricultural abandonment, presence/absenceof cuttingin the previous 100 years, historicalfrequencyof turnoverin ownership, and the distinctionbetween primary(always forested) and secondary (formerlycleared land) woodlands (cf. Peterken1981). Results WORCESTER SETTLEMENT LAND-USE AND VEGETATION HISTORY The historyofland ownership(1738-present) ofthe ProspectHill tractwas compiled fromproprietors' grants, deeds and tax valuation lists. Complete ownership chronologies were compiled for each of the 25 propertylots that comprise the tract. Turnoverrates in ownershipwere calculated on a decadal basis as the percentageof the 25 lots that had been sold out of a family.Land-use history was derivedfromvarious sources. Deeds and sale recordsprovideinformationon lot statusas either wooded, pasture, mowingsor tilled field, and on timber rightsheld on woodlots. For each forest stand in the studyarea the followinginformation had been reconstructed by earlierresearchersat the Harvard Forest: date of clearance, agriculturaluse, timing of abandonment, successional vegetation, and historyof logging(Fisher1921; Raup & Carlson 1941; Spurr1950; Gould 1960). COUNTY - EARLY POST- VEGETATION On a regionalbasis the distributionof forestvegetationin the 1700s closely matchesthat of today. Forest types includingCentral Hardwoods, TransitionHardwoodsand NorthernHardwoods (Fig. 6) axis across are arrangedon a south-eastto north-west WorcesterCountyfollowingthe altitudinalgradient (Figs 1 and 7, Table 1; Westveld 1956). Central Hardwood forestswere dominatedby Quercusspp., Carya glabra, Castanea dentataand Pinus strobus. Other species includedAcer rubrum,Pinus rigida, Betula spp. and Fraxinus americana. Transition Hardwoodforestsincludethesame species,withmore Acer rubrum,Betula lenta and Fraxinusamericana as well as Fagus grandifolia,Tsuga canadensisand occasional Picea spp. and Larix laricina. In the NorthernHardwood forest,Carya, Castanea and Pinus rigida were minor components; additional common species included Acer saccharum,Betula alleghaniensisandB. papyrifera.In addition tothese This content downloaded from 128.103.149.52 on Tue, 3 Sep 2013 16:19:33 PM All use subject to JSTOR Terms and Conditions 759 D.R. Foster Table 1. Forest compositionof the townshipsin WorcesterCounty,Massachusettsin the late 1700s as reportedin the journal of Peter Whitney(1793) and interpretedfromdetrendedcorrespondenceanalysis (Fig. 6). The townsin each foresttypeare listedat the bottomof the table. Modal Value (MV) of abundance refersto Whitney'sdescriptionas (1) occasional, scatteredor rare, and (2) commonor dominant.Frequencyis the percentageof townsin a forestregionin whichthe species occurred Carya ovata Juglanscinerea Platanusoccidentalis Ulmusamericana Sassafrasalbidum Quercusprinus Chamaecyparisthyoides Carya glabra Pinus rigida Castanea dentata Quercusspp. Pinus strobus Acer rubrum Pinus resinosa Betula spp. Fraxinusamericana Fagus grandifolia Tsuga canadensis Picea spp. Larix laricina Fraxinusspp. Betula lenta Acer saccharum Betulapapyrifera Betula alleghaniensis Tilia americana Central Hardwoods TransitionHardwoods Pine and Flood Plains Transition Hardwoods Northern Hardwoods MV MV Frequency MV Frequency MV Frequency 2 2 1 2 1 2 0-50 1-00 0-83 0-66 0-33 0-17 2 2 2 2 1 2 1-00 0-83 1-00 1-00 0-83 0-33 1 1 2 1 1 0-66 0-66 0-17 0-17 0-17 1 0-17 1 2 1 2 1 2 1 2 2 2 2 1 2 2 2 2 1 1 2 2 1 0-04 0-24 0-04 0-04 0 04 0-92 0-48 0-96 1-00 0-76 0-72 0-16 0-80 0-80 0-48 0-40 0-08 0-04 0 04 0-08 0-04 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 0-20 0-60 0-80 1-00 1-00 0-60 0-20 1-00 0-80 0-80 0-80 0-40 0 40 0-40 0-60 1.00 0-40 0-60 0-20 1 2 2 2 2 1 2 1 2 2 Frequency 0-45 1-00 0-45 1-00 1-00 0-91 0-36 0-09 0-27 0 18 Forest region Townships CentralHardwoods Northbridge,Oxford,Southborough, Douglas, Dudley, Mendon, Milford,Northborough, Upton, Uxbridge,Webster,Westborough TransitionHardwoods withPlains Grafton,Harvard,Lancaster,Leominster,Sutton,West Boylston TransitionHardwoods Athol, Barre, Bolton, Charlton,Fitchburg,Hardwick,Holden, Hubbardston,New Braintree,Northbridge,Paxton, Petersham,Phillipston,Princeton,Oakham, Rutland, West Brookfield, Shrewsbury, Spencer, Sterling,Sturbridge,Templeton,Westminster, Worcester NorthernHardwoods Ashburnham,Gardner,Lunenburg,Royalston,Winchendon No information Berlin,Clinton,Leicester major foresttypes,in the easternhalfof the county on floodplainsand sandyoutwashthereoccurreda distinct assemblage of trees including Platanus occidentalis, Ulmus americana, Carya ovata and Juglanscinerea. PETERSHAM FOREST - EARLY VEGETATION AND CHANGE Peter Whitney(1793) described the original vegetation of Petersham as predominantlyQuercus on the uplands and Betula, Fagus grandifolia, Acer rubrum,Fraxinus, Ulmus and Tsuga in the lowlands. Castanea and CaryalJuglanswere noted as increasingfollowingsettlement.Changesin forest cover throughtimeforPetershamparallel those for other townshipsin Worcester County (Black & Wescott 1959), but were more rapid and greater as a whole(Fig. 3). thanforthestateofMassachusetts At the heightof agriculturalactivity(mid 1800s), the Petersham landscape was a mosaic of small fields(indicatedby thepatternof stonewalls;Figs 2c and 5b) and isolated woodlands (Fig. 4). Reforestation proceeded outwardfromthe forestremnants, leaving progressivelysmaller areas of open land along the ridgetops (Fig. 4). One major patternof This content downloaded from 128.103.149.52 on Tue, 3 Sep 2013 16:19:33 PM All use subject to JSTOR Terms and Conditions 760 Land use and vegetationhistory in New England (a) 200 *Rut Rt %% *Gar Roya 4 Pinus rigida Plains *Sut rAsh@ Northern *Hub *Petersham * WBWesmHa WrEa HardwoodsAth *Stu NBr N 150 DHardwoods Win L_nh Lan 0~~~~~~~ WB WBo "Upt PaBar *Pri/Har *te/Pil Ste Cha *Spe G *Gra % % ?l-Worh NBro@ 100 and Flood Plains Transition Hardwoods *oxf *NBo *WBo *Dud/Fit SBo/Uxb * NBri Waro Men/Oako 50 _ Central Hardwoods O I \Mil, cm 2A x * Sas alb (b) 400 200 - Betpap* Oo * Larlar * Pic SPP. * Tsu can Fag gra* Betspp *Ace rubFraspp # * Ace sac * p Ulmame * Jugcin Pla 0cc * Car ova * Pinrig * Fraame Pinres ** Pinstr * Betlut Que spp. *Betlen Bet len 0 *C * r gla den *~~~~~~~0Cas -200 * Cha thy -400 -200 0 Axis1 200 400 of theforestvegetationof Worcester Fig. 6. Sampletownships (a) and species(b) plotsforPeterWhitney's(1793) description County,Massachusettsdisplayedon the firsttwo DECORANA axes. Townshippoints(a) are groupedintomajor vegetation types (NorthernHardwoods, TransitionHardwoods and Central Hardwoods). The Pinus rigida plain and flood plain variantoftheTransitionHardwoodsis indicatedbysquaresymbolsand separatedbya brokenline.Town namesare provided in Table 1. Townshipabbreviationsuse the firstthreeor fourlettersof the name. Species abbreviations(b) are based on the firstthreelettersof the genusand species: Acer rubrum,Acer saccharum,Betula lenta,Betula lutea,Betulapapyrifera, Betula spp., Caryaglabra, Carya ovata, Castanea dentata,Chamaecyparisthyoides,Fagus grandifolia,Fraxinusamericana, Fraxinusspp., Juglanscinerea,Larix laricina,Picea spp., Pinus strobus,Pinus rigida,Pinus resinosa,Platanusoccidentalis, Quercus spp., Sassafrasalbidum, Tsuga canandensisand Ulmusamericana. naturalreforestation includedestablishment ofPinus strobuson abandoned pastures and fields (Fisher 1918). An alternativepatterninvolvedsuccessional hardwood species, including Betula populifolia, Populus spp., Acer rubrumand Prunus spp. Over time, as eitherthe Pinus strobuswas harvestedor the successionalhardwoodsdied, the modernforest of Quercus, Acer rubrum,Pinus strobus,Fraxinus americana,Betulalentaand B. alleghaniensis emerged (Fisher 1933). PROSPECT HILL TRACT Historyof land ownershipand development Historicalchangesin the size and durationof individuals holdings reflectthe type and intensityof land-use and indicate ongoing modificationof the forestlandscape. Duringthe 160-yearperiodbefore acquisitionby Harvard Universityin 1907 each of the 25 lots comprisingthe Prospect Hill tractwas This content downloaded from 128.103.149.52 on Tue, 3 Sep 2013 16:19:33 PM All use subject to JSTOR Terms and Conditions 761 D.R Foster 400 - NorthernHardwoods Ash Gar 0 300300 E Cha .oS W Br Spe Stu < 200 ou Of Mil \BaS 42?00' Roy Win Fit / Gra WBo NBo Har Upt "Plains"' Bol L Lan Bo ardwoods \4Cenral Leo Ste Sut Uxb Men NBri 0 WBoy Wor tDud~~ 100 Hub Phi * * *~~~~~~~~ Wes... Tern Pri Rut Oak Bar Pet * * N Br Hard *HoI Ath * Transition Hardwoods * Pax g 42?10' 42020' 42030' 42040' Latitude North Fig. 7. Location of the townshipsof WorcesterCounty, Massachusettson altitudinaland latitudinalgradients.Major vegetationaltypesare indicated.CentralHardwoodsvegetationis foundpredominantly at low altitudein thesouth-eastern cornerof the county,whereasthe townshipsin the elevated north-western area are characterizedby NorthernHardwood forests.Townshipsin the Pinus rigidaand floodplain subtypeof TransitionHardwoods are markedby square symbols. Townshipabbreviationsuse the firstthreeor fourlettersof the name. Proprietors'grants 100100~~~~~~~~~~ .-. III IV Harvard Forest-> Jonathan Sanderson firstpurchase 8080 / / \Adonai Shomo 60 o -j Sandersonprobate 4020 - Lots transferredto descendants I 1740 *''I 1760 I 1800 I I I 1840 " 1880 p I 1920 I_ 1960 Date Fig. 8. Historicalchanges in ownershipturnoverforthe ProspectHill tract,HarvardForest. The graphdepictsthe perto directdescendants centageof lots in the tractthatwere sold in the precedingdecade. The percentageof lots transferred (generallysons) is indicatedby the brokenline and largersolid dots. sold an averageof 13 times.From 1733to 1770turnover exceeded 80% on a decadal basis as absentee ownersspeculatedon land grantsand sold theirlots to the firstsettlers(Fig. 8). During the commercial agriculturalperiod through the mid 1800s, the lowest level of turnoveroccurred (10-40% on a decadal basis) as families retained their land, worked it intensivelyand graduallyexpanded their holdings throughacquisition of adjoining parcels (Fig. 9). After1840 and coincidentwiththe beginningof farmlandabandonment,ownershipturnover increased markedly; owners often held parcels for only 4-12 years between sales. A temporary decline in land sales occurredduringthe economically depressed period of the American Civil War (1861-65; Coolidge 1948). This content downloaded from 128.103.149.52 on Tue, 3 Sep 2013 16:19:33 PM All use subject to JSTOR Terms and Conditions 762 Land use and vegetationhistory in New England 1780 1770 1760 N 3 2 1810 1800 1790 :. t; ~~~~~~~~~~~~~~~~~~~~~~~..... /oi ~ . Sa~~~~nderson McClellan Pierce 1870 ~~~~~~1860 Mann Pierce Ward /Spooner ~~~~~~Spooner Ado(o Shomo nai Shomo Adonai Hodges Marsh / Briggs eta!. Spoonerd r Sanderson 1890 1880 X Sanderson Ward anderson Ward Pierce Ward . rson Farnsworth McClellan 1840 1830 anderson Pierce 1850 aswo arn Farnsworth McClellan Wcllard 1820 Farnsworth McClellan Goddard Farnsworth McClellan Goddard f Willson Knapp Chandler / 4 1900 Adonai Shomo ~Russell Marsh M h Leonard Stone - Fig. 9. Ownershipmaps at 10-yearintervalsfor the period 1760-1900 on the ProspectHill tract,Harvard Forest. The propertiesof the threelargestlandownersforeach decade are indicatedas (1) black, (2) diagonal lines, (3) shaded. The names of theownersof theseareas are listedto thebottomrightof themap in descendingorderof propertyarea. Growth in the size of the largerfarmsis indicatedby the Sandersonfarmfrom1800 until1840. This content downloaded from 128.103.149.52 on Tue, 3 Sep 2013 16:19:33 PM All use subject to JSTOR Terms and Conditions 763 D.R. Foster Farm development Agricultural historyon theProspectHill tractclosely paralleled that of the township.Forest clearance commenced c. 1750, proceeded at a slow pace through 1780, and abruptlyincreased thereafter until 1840-50 when open land peaked at nearly 80% (Fig. 3; Fisher 1933; Raup & Carlson 1941; HF Archives). During the period 1820-80 there was relativestabilityin the patternof open land and forest. The geographic distributionof land-use types relatescloselyto the physiographic and soil characteristicsof the area, and to the distancefrommajor roads and dwellings(Table 2; Figs 2b and 10). The earliest areas to be settled and developed were adjacent to roads and houses. Nearly 50% of this accessible land was tilled (Fig. 10), whereas across the entire tract approximately70% of the area was cleared for pasture, 17% was tilled, 13% was wooded and slightly morethan1% was continuously marsh (Table 2). Permanentwoodland comprised threewooded swampsin the centreof the tractconnectedalong poorlydrainedseepages and adjoining steep or rockysites (Figs 2b and 10). Tilled land primarilyoccupied well-drained and imperfectly drainedsoils whereaspasturecovered the broadest range of drainage conditionsincludingall of the verywell drained soils, 75% of the poorlydrained sites, and nearly70% of the well and imperfectly drainedsoils (Table 3). The processof closelyfitting tillageareas to soil drainageled to irregularly shaped and small fields(Figs 2c and 10). Logginghistory The history of logging includes four periods: (i) land clearance; (ii) woodlot cuttingduringthe agriculturalperiod; (iii) commercialharvestingof old-fieldPinus strobusand old woodlots(1885-95); and (iv) Harvard Forest management. Different objectives on the part of the landowners led to varyingeffectsduring these periods. Once agriculturallandwas cleared,deeds fromthenineteenthcenturyindicatethat the remnantwoodlands were a valuable source of Castanea and Tsuga for fuel wood, timberand tan bark. Forest reconstructions indicatethatwoodlotswere cut repeatedlyforpolesized trees afterthe initial removal of the virgin timberin the mid to late 1700s (E.P. Stephens & S. Spurr,HF Archives;Fig. 11). I N > Pasture and Mowing \/tdWoodlot ? Tilled Land Open Marsh Fig. 10. Land-use historymap forthe ProspectHill tract, HarvardForestdepictingpastureand mowing,woodland, tillageand marshat the timeof mostintensiveland-usein the mid 1800s. Althoughland use changedover time,this map showsthecontinuouswoodlandarea and all sitesthat were ever tilled.ModifiedfromSpurr(1950) and Harvard Forest Archives. of the ProspectHill tract,Harvard Forest. Land-use typesdescribe Table 2. Soil drainageand land-use characteristics the primaryuse of the land duringthe heightof agriculturalactivityin the mid 1800s (Spurr 1950; HF Archives) with areas givenin hectares.In the modernlandscape all of the area is forested.All values were calculatedon a Geographic Information Systemand are roundedto the nearestinteger Land use (ha) Soils Pasture Tilled Woodlot Marsh Totals Very well drained Well drained drained Imperfectly Poorlydrained Verypoorlydrained 14 150 27 45 3 49 9 2 - 14 4 13 14 4 14 213 40 60 21 Totals 239 60 45 4 This content downloaded from 128.103.149.52 on Tue, 3 Sep 2013 16:19:33 PM All use subject to JSTOR Terms and Conditions 764 Land use and vegetationhistory in New England axes and species axes in the canonical variables,environmental Table 3. Weightedcorrelationmatrixforenvironmental correspondenceanalysisof 1937 data fromthe ProspectHill tract,HarvardForest. Environmentalvariablesincludestand woodland, cuttinghistory,and abandonmentdate. primary/secondary age, soil drainage,slope position,pasture/tillage, Env Ax x representsthe sample scores on the xth ordinationaxis, whichare linear combinationsof the environmental variables. Spec Ax x representsthe sample scores on the xth ordinationaxis whichare derivedfromthe species scores by weightedaveraging Age Soil Slope Pasture Primary Cut old field Abandonmentdate Env Ax 1 Env Ax 2 Spec Ax 1 Spec Ax 2 Age Soil Slope Pasture Primary Cut old field Abandonment date 1-00 0-21 0-28 -0-21 0-47 -0-31 0-47 0-79 0-37 065 0-25 1.00 0-44 -0-16 0-34 0.01 0-34 0-54 -0-68 045 -0-46 1.00 -0-34 0-53 -0-17 0-49 0-57 -0-51 047 -0 34 1-00 -0-68 0-31 -0-46 -0-46 -0-08 -039 -0-05 1.00 -0-29 0-84 0-85 0-02 0-71 0.01 1.00 -0-01 -0-23 -0-25 -0-19 -0-17 1-00 0-84 0 09 0-70 0-06 From 1885 to 1895, essentiallyall of the merchantabletimberon the ProspectHill tractwas cut often followed by burningof the slash (Fig. 11) (Fisher 1921; HF Archives).This timbercame from woodlots that were cut for Tsuga, Castanea and Quercus and from former pastures abandoned between 1845 and 1865 and then dominated by Pinus strobus. j vegetationdevelopment Post-agricultural Followingthe heightof forestclearance (1820-80), reduced agriculturalactivityresulted in rapid reforestation (Fisher1921;HF Archives).Today, essentiallythe entirestudyarea is covered by maturing forest(Fig. 12; Gould 1960). The modern forest historyis best understoodby examiningthe pattern of land-abandonmentand subsequent vegetation development. Patternand rateof land abandonment On theProspectHill tractand in centralNew England in general, farm abandonment and reforestation commenced slowly in the mid nineteenthcentury and subsequentlyincreasedgreatlyinto the 1900s. morethan25% of the ProspectHill By 1880 slightly tractwas forested,whereasthisfigureincreasedto 85% by 1937(Figs 3 and 12). The geographicpattern of field abandonmentwas complex, controlledin part by an individualfarmer'seconomic situation and land holdings.However, the generalprocess is consistent:abandonmentfirstof poorlydrainedand low-lyingpastureareas adjacentto the intactwoodland, followed later by well-drainedand arable lands (Fig. 12). From 1840 to 1875 only pastures were abandoned, whereasthe firsttilledfieldswere abandoned in the late 1870s. By 1880 the forested 1830 4 2 F ^ ~~~ ~~~18 70"a.' 1881~~~~~88 i Forest <20 years old In 1907 Open land in 1908 Fig.11. Cutting history oftheProspect Hilltract, Harvard Forestin the periodbeforeHarvardForestownership (1907).Datesinwhiteareasindicate theyear(s)ofcutting. Shaded areas werein immature forest(<20 yearsold) and blackareaswerein openfieldor scrubin 1907.Data reconwerederivedfromhistorical maps,notesandforest storedin theHarvardForestArchives. Muchof structions thewhiteareacontained merchantable timber in 1907. area formeda continuousstand withsmall pasture in-holdings(Fig. 12). Open areas were limitedto 25% of the area in 1900,and followingthe transfer of the land to Harvard Forest,the remainingfields were abandoned or plantedwithconifers. Vegetationdevelopment fromfarmabandonment to present Maps and forest-wide inventories fromthetwentieth This content downloaded from 128.103.149.52 on Tue, 3 Sep 2013 16:19:33 PM All use subject to JSTOR Terms and Conditions 765 D.R. Foster 1840 44 1880-99 1840-59 1860-79 1900-19 1920-present ~~0 ~~~ 500 m Fig. 12. The historyof land abandonmentforthe ProspectHill tract,HarvardForest at 20-yearintervalsstartingat the peak of land clearance in 1840. Areas in forestduringthe previous20-yearperiod are depictedin black. For each period abandoned pastureland is shaded, whereasabandoned tilledfieldsare indicatedby diagonallines. Data werederivedfrom historicalreconstructions storedin Harvard ForestArchives. century(in 1908, 1937 and 1986 - HF Archives) depictstriking relationshipsbetweenvegetationand land-use patternsand dynamicsin the vegetation resultingfrom forest development, management and the 1938 hurricane.In 1908 Picea forestsand the limitedarea of Tsuga forestwere confinedto formerwoodlots (Fig. 13). One-thirdof the former woodlot area was classified as hardwood forest (Quercus spp., Acer rubrumand Betula spp.) with scattered Tsuga understorey. Former pastures were forestedwithPinus (Pinus strobusand Pinus strobus-Tsuga canadensis forest) or successional stands of Betula populifolia, Acer rubrum and Populus spp. Approximately40% of this pasturehardwoodarea had supportedPinus strobus,which was cut in the 1890s. Only 21% of the one-time pasture land was still open in 1908, whereas 53% of the tilledfieldswas open. Major changes from 1908 to 1937 occurredas a resultof standdevelopmentand silviculture (Fig. 13). Nearly 100ha of conifer plantationswere established on fields(42%), clear-cutPinus strobusland (28%) and successional hardwood areas (12%). The majorityof the plantations(81%) were establishedon well-drainedsoils; approximately75% on formerpastures and 25% on tilled fields.The mortalityof Castanea in 1913-16 due to blight (Cryphonectriaparasitica; Kittredge 1913; Spurr 1950) led to the naturalconversionof formerCastanea stands to Tsuga (84%) or hardwood (16%) forest.Overall, hardwoodforestwas reduced from 135ha to 75 ha between 1908 and 1937. However, stands of long-livedspecies (Quercus, Betula lenta, B. alleghaniensis,Fraxinus, Fagus and Acer saccharum) increased from 40 ha to 56 ha, whereas successional hardwood forest (Betula populifolia, Populus and Prunus) decreased greatly (95 ha to 19ha; McKinnon, Hyde & Cline 1935). The reduction in successional species resulted from clearingfor plantationsand selective management for longer-livedspecies (Cline & Lockhard 1925; Gould 1960). Increases in Tsuga (<1ha to 16ha) and Pinus strobus-Tsuga canadensis (12 to 27ha) forestsfrom1908to 1937were largelydue to developmentof foreststhathad a substantialcomponent of Tsuga in the understoreyand perhaps due to a reductionin post-loggingburning(Marshall 1927; Spurr 1956). Withina year of completionof the 1937 forest survey,the forestlandscape was completelyaltered This content downloaded from 128.103.149.52 on Tue, 3 Sep 2013 16:19:33 PM All use subject to JSTOR Terms and Conditions Landuse Soil drainage ?S - .19% * Verypoor 66?b Poor t75 - - (4 ha) 'Marsh/Swamp -88%< Woodlot(45 ha) 831% Verygood 100% Imperfect67% ------------22%Y------------------1%- - 6% Pasture(239 ha) r62% Tle(6h) Ild(0a GoodF .23% -- Landuse 1908 Foresttype Marsh/Swamp(4 ha) Lt56%- 3- Tilled3(XOha) ( ** 37% Open (87 ha) Acer rubrum(33 ha) a%ia 2A% Pasture( 239 ha) -2% 79,%<- Pinusstrobus(64 ha) Castanea (4 ha) 8?0.% - Hardwoods(135 ha) Pinusstrobus-Tsuga( 12ha) 3' Woodlot (45 ha) 4-23X___Tsuga(<lha) -23% 2%*--__ ---- by the 1938 hurricane(Foster & Boose 1992). Coincidentally,the management objectives of the Harvard Forest shifted to favour natural stand development rather than timber production of conifers(Gould 1960). The changes wroughtby natural disturbance and forest development are reflectedin major changes in forest composition from 1937 to 1986 (Fig. 13). Overall there was a major increasein hardwoodstands(75 ha to 188ha) resultingfrom preferentialhurricane damage to plantations(reduced from96 to 35ha) and Pinus strobusforests(56ha to 25 ha), cuttingof conifers, and old-fieldsuccession (Spurr 1956; Foster 1988; Foster & Boose 1992). Open areas were reduced by65%, Acer rubrumforestdoubled (17 ha to 35 ha) and Tsuga forestscontinued to increase through standdevelopment.Nearly35% of the Tsuga stands developed following hurricane damage, cutting and natural succession in hardwood stands, 41% fromhurricane-damagedPinus strobusand Pinus strobus-Tsuga canadensis stands and 20% from hardwood stands, includingAcer rubrumswamps (HF Archives). 96%-m- Tsuga( <I ha) ---'87%- Plcea(12 ha) LAND-USE HISTORY AND FOREST COMPOSITION 1908 Vegetation 1937 Vegetation 42%- Pinusstrobus(64 ha) 39%- - 11% Open (87 ha) -28%.sI s4 * --. .7 ji% Open (49 ha) ~~~~~~11% Acer rubrum(33 ha) 1-216%49*l rubrum 349%3hI A2ce% 14% - 2 = Hardwoods(135 ha) V44% PinusstrobtisTsuga (12ha) Plantation(96 ha) s Pinus strobus(56 ha) ..- Acer rubrum(17ha) 78%-*4 Hardwoods(75 ha) ------------------- = 927%.74.%r~k Plnusstrobus-Tsuga(27 ha) Tsuga(< Iha) i 93%.3% Castanea (4 ha) ,-84% 21%-1 Tsuga(16 ha) Plcea (I 2 ha) 87%...................................-96% .. - P/cea(I I ha) 1986 Vegetation 1937Vegetation - Plantation (96 ha) t33%- 90%-)-i 1.68%t%~-~~ Open(49ha) Plantation (35 ha) Open(16ha) '9T~ 2%-p.- Hardwoods(188ha) (75 ha)t8>X%_,,,,-685% Haurdwoods Plnus strobus (56 ha) - ----- 70% -15%----------- 4 r r 5ce Tsuga(lG6ha)| 365%. Pinus strobus-Tsuga(27 ha) Pccea (1I ha)' 83%_. 17.- Pnus strobus (25 ha) -- ;Acer rubrum(35 ha) bu(17h)t%~~'~---******34%..<j - 55% . .,, ,_,. Canonicalcorrespondenceanalysisofthe 1937forest data identifiesconsiderablevariationin stand compositionthatmay be explainedusinga combination of historicaland site factors.A minimalset of variables thatbest explainsthe data was identified using theoptionofforwardselectionofvariableswitheach variabletestedand an overalltestapplieldby means test(terBraak 1986). of a Monte Carlo permutation Site factorsselectedincludedsoil drainageand slope position, whereas important historical variables included: primary/secondarywoodland, cutting history,abandonmentdate, and pasture/cultivation. This model was significant at the0-01level. The first two CCA axes (Fig. 14) account for 62% of the species-environmentrelations. Samples separate out clearly along the landuse and soil-drainagegradients(Fig. 15). Primary woodlands are concentratedon the intermediateto poorlydrainedsites,whereas secondarywoodlands occurmorewidelyon all buttheverypoorlydrained soils(Fig. 15). There are strongcorrelationsbetween variables,e.g. between manyof the environmental primarywoodland and abandonmentdate, forest 5%::I Tsuga(3Oha ) Hardwoods(188ha) Fig. 13. Transitiondiagrams depictingrelationshipsbetweenland-use,soilsand vegetationin 1908,1937and 1986. The areal extentof each categoryor vegetationtype is indicatednextto it. Percentagesreferto the contribution to percentagesof the adjacentcategoryor typeand do not equal 100% forsome typesas onlymajor relationshipsare depictedforclarity. This content downloaded from 128.103.149.52 on Tue, 3 Sep 2013 16:19:33 PM All use subject to JSTOR Terms and Conditions 767 D.R. Foster age and slope position,and between slope and soil drainage(Table 3). These correlationsare apparent on the species and stand biplots as axes for the environmentalvariables lie closely and parallel to each otherand can be used to interpretstanddistribution.The distribution ofprimarywoodlandstands along the firstaxis is largelyexplainedby variation in age of the forestas controlledby cuttinghistory. A numberoftreeand shrubspecieshave an affinity for the primarywoodland and swamp sites; these include: Tsuga canadensis, Picea rubens,Fraxinus nigra,Nyssasylvatica,Viburnumalnifolium,Kalmia latifolia,Hamamelis virginiana,llex verticillata and Nemopanthusmucronata(Fig. 14). The Tsuga and swamp forestson primarysites generallysupport littleherbaceousgrowthdue to deep shade; consequently,the herbdistribution on the CCA does not showanyspecieswithan affinity forprimaryforests. The secondary forestsare characterizedby early successionalspeciesand speciesofricher,driersites. Pru pen Trees 0.3 Age Cor cor. Til ame. sac AceAcec -Fra ame Bet len Fag gra . Que . t Ulm ame Pruser -0.3 Fig. 14. Species and environmental variable bi-ploton the firsttwo CCA axes for 1937 vegetationdata fromthe ProspectHill tract, Harvard Forest. Species plots are separated bygrowthformintotrees,shrubsand groundcover. Species abbreviationsare based on the firstthree lettersof the genus and species: Trees - Acer rubrum,Acersaccharum,Acer spicatum,Betula lenta, Betula lutea, Betula papyrifera, Betula populifolia, Corylus cornuta, Fagus grandifolia, Fraxinus americana, Fraxinus nigra, Nyssa sylvatica, Picea rubens,Pinus strobus,Plantationspp., Populus tremuloides,Prunus pensylvanica, Prunusserotina,Quercusrubra,Quercusspp. Tilia americana, Tsuga canadensis, Ulmus americana;Shrubs - Azalea spp., Crataegus spp., Hamamelis virginiana, Ilex verticillata, Kalmia latifolia,Nemopanthusmucronata, Ostrya virginiana, Rubus allegheniensis, Rubus hispida, Vaccinium corymbosum, Vacciniumpensylvanicum,Viburnumacerifolium, Viburnum alnifolium, Viburnum cassinoides, Viburnumdentatum;Herbs Aralia nudicaulis,Asterspp., Athyrium filixfemina, Cypripedium acaule, Clintonia borealis, Coptis groenlandicum, Cornus canadensis, Dennstaedtia punctilobula, Fragaria virginiana,Gaultheriaprocumbens, Gramineae, Lycopodium complanatum, Lycopodium obscurum, Lycopodium lucidulum, Maianthemumcanadense, Mitchella repens,Monotropa uniflora,Onoclea sensibilis, Osmunda cinnamomea,Parthenocissus quinquefolia, Pteridiumaquilinum, Pyrola rotundifolia, Rhus radicans, Smilacina racemosa, Solidago spp., Sphagnum spp., Trientalisborealis,Viola spp. Environmental variables are labelled on the tree plot, with Pop tre !. Oron Abandonment Date Pic rub * . Bet lut Ace rub Ace spi . Pasture Primary 0.5 Bet poP . Fra nig Nys syl. Slope Soil --0.4 -0.2 Shrubs Craspp Vibaln * Kal lat Vib ace _0 I I Aza spp . I Hamvir Rub all . Vac cor. Vibden I* le vir * Nem muc o -0.4 Herbs T0.2 DCypaca Lyccom.jLyc luc bor Pteaqu* Tri Mon Mal Mnuni Smi-rai Ican Gra spp Pyrrot . Lycobs Gau pro. Rhurad -0r3 Sol spp. .Astsppprqui CorCan Ara nud Athfil . Osm cin Vio Spp * Ono sen * Copgro the centroids of nominal variables (primary/ secondary woodland, pasture/cultivation, cuttinghistory)indicatedby a large square, and ordinal variables (age, abandonment date, soil drainage, and slope position) indicatedby arrows. pop ;Bet * cut..s Pl a p Tsu can . S * Sphapp -0.4 This content downloaded from 128.103.149.52 on Tue, 3 Sep 2013 16:19:33 PM All use subject to JSTOR Terms and Conditions o!s p Secondaryforests 768 Land use and vegetationhistory in New England forests Primary porydrie wel drainy Very /~~~ Porydri ~~ Fig 1. amle isriutonfor25 pot o th frs CA xes Te ivsio btwenprmar ad ecndry orst aon Axs1 siniatdbytevetca roe ln. oldriag lass r eprte y oidwvylns.Otles ihi h drainage~Alse ar niae ydfeetsmos eog fa sml otenx re oltp ti ersneya open~~~~~~ dotifi eog ~ ~~~ otenx ose ol yei sidctdb Discussion One major conclusionderived fromthis historical analysis is that at any scale - forest,township, countyor state- land-usepracticesand theresulting vegetation patterns have changed continuously (Bradshaw & Miller 1988; Fosteret al. 1992). Over the past 250 years there have been few periods of stabilityin whicheven the extentof forestvs. open land was relativelyconstant (Fig. 3). Population density, the agricultural,forestryand industrial practices,and the geographicalpatternof land use were all highlyvariable. The changingqualityand intensityof human activityresultedin the dynamic vegetationcharacteristic of thisperiod. LAND-USE HISTORY Across centralNew England there has been great similarityin the regional pattern of land use in terms of the extent and timingof deforestation, major agriculturaluses, and the historyof farm abandonment and reforestation(Bogart 1948; Gates 1978; Taylor 1982; Garrison 1985). The ProspectHill tractof the Harvard Forest is clearly of thetownshipof Petershamin these representative characteristics (Figs 3, 4 and 12). WorcesterCounty, in general,was more agriculturally orientatedthan westernparts of the state, and thus the rate and rage extentof forestclearingin Petershamare greater than state averages. However, temporal shiftsin land use followedthesame generalmode throughout upland areas: land clearance and gradual depletion of thetimberresourcewas followedby low-intensity agriculture,commercial agricultureand industry, and subsequently by population migration and farmabandonment(Bidwell 1916; Garrison 1985; Williams1989). In the modernperiod,utilizationof the land fornaturalresourcesis at a historicallow, but pressurefor housingand commercialdevelopmentis leadingto a minorsecond wave of clearance, and a concernforforestpreservation(Anonymous 1940; Black & Wescott 1959; Bickford& Dyman 1990). In thispattern,Petershamparallelsthetrends of manyupland townships(MacConnell 1975). The land-usepatternthatemergeson examination of the Prospect Hill tract is exceedinglycomplex and variable in scale and intensityover time. Each of the individualparcels comprisingthe tractwas ownedbyan averageof 13 individualsin the160-year period. These owners differedin the size of their holdings,durationofownership,and undoubtedlyin theireconomicand social perspective(e.g. Garrison 1985). Althoughthe details vary,each foreststand was cut repeatedly,mostwere cleared, and all were subjected to varying agriculturaluse, including grazing (Graves & Fisher 1903; Barraclough & Gould 1955). This content downloaded from 128.103.149.52 on Tue, 3 Sep 2013 16:19:33 PM All use subject to JSTOR Terms and Conditions 769 D.R. Foster of the geographicpatternof In the determination land use there were strong interactionsbetween environment,spatial relationships and land-use quality. Each land-use activitytriggeredvarious vegetationalchangesthataffectedsubsequentlanduse was use decisionsup to thepresent.Agricultural stronglycontrolledby site conditions(undoubtedly includingthe original vegetation; cf. Lord 1973) and distancefromthe road or farm.Sites adjacent to farmsand roads were cleared first,used most intensivelyand abandoned last (Figs 2c and 10). Soil moisturestronglycontrolleduse, withwet sites used as woodlots, mesic sites tilled for cultivation and extremelydryor moistsitespastured(Table 2). WithinPetershama similaroverall patternof use based on distance and soil conditionsappears to hold (Fig. 4) (Raup & Carlson 1941). The quality and timingof land-use determined the subsequent vegetation,which in turn affects later human activityand the impactof naturaldisturbance processes. For example, the extensive loggingenterprisein the late 1800s and early 1900s was controlledby the timingand placementof field abandonmentin the mid 1800s. Similarly,decisions concerningthe placement of conifer plantations were determinedby the distribution of open fields or early successional vegetation(formerfields) in theperiod 1910-40. In termsof naturalprocesses,it is clear that the effectof the 1938 hurricanewas controlledby the structure,compositionand spatial distribution of the vegetationthatresultedfrom200 yearsof humanactivity(Foster & Boose 1992). The Pinus strobusand plantationson formerfieldswere particularlysusceptible to wind damage and are largelyresponsibleforthegreatextentof damage to the landscape of centralNew England (Rowlands 1941). VEGETATION RESPONSE TO LAND USE Evaluation of the effectof cumulativeland use on the vegetationis stronglyscale-dependent.Across the region (i.e. Worcester County) the effectis subtle and suggeststhat the vegetationis resilient to disturbance(Raup 1966). Species assemblages sorted out in the 1700s along climatic gradients definedby latitude and altitude much as they do today (Westveld 1956). However, shifts in the a general abundanceofspeciesare notable,primarily increase in successionaland sproutingspecies (e.g. Betulapopulifolia,Acer rubrum,Quercusspp.) and declinein long-livedtolerantspecies (Fagus, Tsuga, Acer saccharum) (Cook 1917; Dickson & McAfee 1988). Pathogenshave also reduced the abundance of Castanea, Fagus and Ulmus (Gravatt & Parker 1949; Hirt 1956; Karnosky1979; Paillet 1982). At a local scale withina forestthe effectsof land use are compelling(Foster & Zebryk 1993). The pattern of vegetation is clearly abrupt and dis- continuousin compositionand structure;gradients are steep. Structurallythe forestsare young and even-aged. In additionto structuralcharacteristics, major compositionaltrendscan be tied to land use. Picea rubensand Tsuga are largelyconfinedto the areas of permanentwoodlot(Fig. 13; cf.Smith1950; Kelty 1984), and a few tree and shrubspecies (e.g. Viburnumalnifolium,Kalmia latifolia,Hamamelis virginiana, llex verticillata,Fraxinus nigra, Nyssa sylvatica)are associated withthese primaryforests (Fig. 14). In contrast,pastureareas are dominated by Pinus strobus,Pinus strobus-Tsuga canadensis, Castaneaand poor hardwoodforestsof Betulapopulifoliaand Acer rubrum(Figs 13 and 14). compositional Duringthe periodof reforestation, changesin thevegetationhave accompaniedchanges in the totalcover, and increasesin mean heightand age. Until the 1938 hurricane,hardwoodsdeclined as a result of increases in Tsuga from advanced regeneration,conversionof successionalhardwood foreststo Pinus strobus,and silviculturalmanagement of mixed stands to increase Pinus strobus and Tsuga. Followingthe hurricane,whichgreatly damaged plantations and Pinus strobus stands, and subsequent salvage logging, hardwood cover increased greatly.Hardwoods continuedto establish in manyopen areas as active managementfor conifersceased. One major trendacross thisperiod is the increasein Tsuga as a major cover type;20% of the Tsuga forestderives from succession and advance regenerationin hardwood stands, 35% from hurricane-damagedhardwoods and 41% fromhurricane-damagedPinus strobus and Pinus strobus-Tsuga canadensisforest. One major questionnot addressedby the current study but warrantingadditional investigationis the longevityof these land-use impacts in terms of forest communitycomposition and structure. Analysis of the 1937 vegetation defines striking relationshipsbetween composition and historical variables, including primary/secondaryforest, age, and timingof abandonment,and site factors (cf. Whitney& Foster1988). Analysisof themodern 55 yearslaterwillenable vegetation,approximately us to see if additionaltime forforestdevelopment and species dispersal and the impact of the 1938 hurricanehave reducedthe imprintof the historyof land-useon vegetationpatterns. Conclusion The continuallydynamicnature of the vegetation patternin centralNew England is one of the most landscape. remarkableaspectsofthepost-settlement occurredin the regional A completetransformation and local landscape from1770 to the presentas the farmedand subsequently was deforested, countryside reforested.Withina landscape such as that of the Harvard Forest, thishistoryof use was dependent This content downloaded from 128.103.149.52 on Tue, 3 Sep 2013 16:19:33 PM All use subject to JSTOR Terms and Conditions 770 Land use and vegetationhistory in New England on originalsite factorsand has greatlyaffectedsubsequentvegetationcharacteristics. These effectsare long-lastingas they continue to control the way in which humans use sites and the ways in which naturalprocesses affectthem. The ramifications of this historyin terms of contemporaryecological processesare too greatto be dismissedby modernday ecologists. Acknowledgments Ernest M. Gould, Jrand Hugh M. Raup provided valuable insightinto the land-usehistoryof central New England and the HarvardForest. G. Whitney on historicalsources offeredinnumerable suggestions and references.A. Lezberg assistedwithfieldwork and GIS analysis, and G. Carlton, M. Fluet, F. Gerhardt,S. Hamburg,C. Mabry,G. Motzkin,W. Niering,R. Peet, G. Peterkenand P. Schoonmaker offeredconstructivecommentson the manuscript, typedby D. Smith.This studyis part of the Long Term Ecological Research Programat the Harvard Forest and was supported by the U.S. National Science Foundationand A.W. Mellon Foundation. 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