The Past and Present Vegetation of High Point State Park, New Jersey Author(s): William A. Niering Source: Ecological Monographs, Vol. 23, No. 2 (Apr., 1953), pp. 127-148 Published by: Ecological Society of America Stable URL: http://www.jstor.org/stable/1948517 . Accessed: 18/04/2013 11:13 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
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VEGETATION OF HIGH POINT STATE PARK, NEW JERSEY
THE PAST AND PRESENT
WILLIAM
A.
NIERING'
Botany Department, Rutgers, the State Universityof New Jersey New Brunswick, N. J.
TABLE OF CONTENTS PAGE INTRODUCTION
.
127
............................
METHODS
..........
..................
RESULTS
.........
...................
Xerarch Succession ......................... Hydrareh Succession ........................ Abandoned Fields ........................... ................. Pitch Pine-Oak Communities ..................... Chestnut Oak Community
129 130
130 131 134 135 136
INTRODUCTION The objectiveof this paper is to describeand correlate the past and presentvegetationalpatternsand point out future successional trends in the plant of High Point State Park. The descripcommunities tions of vegetationare intendedto be suitable for generaluse as well as to contributeto thevegetational New Jersey. High Point knowledgeof northwestern State Park was selected for the study since no detailed investigationshad been made in the region. Furthermore,it representeda delimitedarea which was relativelyfreefromdisturbancesince its acquisition as a State Park. The park is located in Sussex County in northwesternNew Jersey. It is an area of 10,935 acres lying just south of the New York State line and saddlingthe crestof the rockyKittatinnyMountains for a distanceof 8 mi. (Fig. 1). Its southernboundary is contiguouswithStokes State Forest. In width it ranges from2 to 4 mi. dependingupon various irregularitiesof the boundary. The area has not long been a State Park. It was givento the State of New Jerseyby the late Colonel and Mrs. AnthonyKuser in 1923. They had built a 220 ft. monumentat the highestpoint on the ridge which is also the highest elevation in New Jersey (Fig. 1). The monument affordsan excellentperspectiveof the park. During aspect the peak of autumncolorationthe differential 'At present with the Department ollege, New London, Connecticut.
of Botany,
Connecticut
PAGE
White Oak-Hickory Community .............. Mixed Oak-Hardwoods Community ........... Northern Hardwoods Communities ............. Pollen Analysis ............................. DISCUSSION .................................. SUMMARY LITERATURE
.................................. CITED
..............................
138 139 140 141 142 146
147
of the various communitiesis pronounced. Light green pitch pine characterizesthe rock outcropsof the ridge top, witha colorfuloak forestpredominating on the slopes. Dark greenconifersare conspicuous in the bogs and swampy areas surroundingthe lakes (Fig. 2). The high series of ridges in the park owe their presence to the geological structureof the region. Physiographically, this part of the State lies in the Ridge-ValleyProvince. The rocks are of Paleozoic age. The limestones, shales,sandstones,and conglomerates-weresubjected to normaland thrustfaulting as well as complexfoldingduringthe latterpart of the Paleozoic. In Tertiarytime,the land was base leveled formingthe Schooley peneplain (Kummel 1940). With the upliftof the Schooleypeneplainin late Tertiary time, the streams were rejuvenated. The harderstrata of sandstoneand conglomerateresisted rapid erosionand today persistas ridgeswhile the less resistantshales and limestonesformadjacent valleys. East of the Kittatinnyridge, shales and limestonesunderliethe Kittatinnyvalley. The ridge is double crested,the easternpart formingthe major ridge (Fig. 2, 3). It consistsof Shawangunkconglomeratewhich is primarilyquartziticwith small whitequartz and slate pebblesembeddedin a siliceous matrix. To the east the ridge drops offas a steep, and in places, precipitousescarpment. Toward its base the Martinsburgshale outcrops,its beds dipping northwestunder the conglomerate(Salisbury 1902).
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128
WILLIAM
A.
PITCH PINE-OAK CHESTNUT OAK SnJ WHITEOAK-HICKORY MESIC COMMUNITIES Eig WE]_ AND 80S COMMUNITIES SWAMP CJ ABANDONED FIELDS
FiG. 1. Vegetational map of I-ugh Point State Park. Locations of stands (1-19) studied in detail are. designated by solid dot adjacent to the number. Other areas studied in less detail are designated by numbers 20-25. Transects A, B, C, D, E, are indicated by solid lines. Stippled areas in eastern sector of park representpitch pine-scrub oak communityon the conglomerate ridge top. In general these areas correspondto the high rocky outcrops. In most cases, the streams can be located from the pattern of the mesic communities. The mesic communitiesinclude the mixed oak-hardwoodsand northern hardwoods. Lake3 are indicated by wavy lines (2024). In the east central sector of the park is the Sussex reservoir (Lake Rutherford). Heavy irregular lines indicate roads. High Point Monument is located at the terminus of the road south of stand 2. Lower lrightoutline map of New Jersey with Sussex County and park located.
However, on the ridge top near the park entrance a small lobe of shale extends northward to Lake Marcia. The lower western ridge crest is developed upon the High Falls formation consisting of red, green, and olive-colored sandstone and shale with interspersed beds of conglomerate. The topography is less rugged than where the Shawangunk occurs. Maximum elevations on the two geological formations are strikingly different. The Shawangunk ridge where the High Point Monument is located rises 1803 ft. above mean sea level (Fig. 1). In contrast the highest ridge on the High Falls is only 1480 ft. During the Pleistocene epoch, the area was glaciated. At least two and possibly three ice sheets advanced southward covering the northern part of the Each modified the topogState (Kummel 1940). raphy by both scouring the highest ridges and elsewhere leaving in its retreat extensive glacial deposits. The pronounced glacial features of the area are due to the most recent glaciation, the Wisconsin-specifi-
NIERING
Ecological
Monographs 2 3, INo. 2
'Vol.
cally the Cary substage. the final Mankato advance terminatingfurther north (P. MacClintock, personal comnmunication).The terminal moraines of the Wisconsinextendas far south as a line extending across the State throughPerth Amboy,Morristown, Hackettstownand Belvidere. Glacial striae on the scoured conglomerateoutcrops indicate that the ice sheetmovedacross the area of the park in a direction. The topographyinfluenced southwesterly the distributionof the glacial drift. It is estimated to reach 10 ft. or more in depth on the High Falls whilealong the crestof the Shawangunkridgeit does not exceed 2-3 ft. (Salisbury 1902). Over the latter it is sporadic and interspersedwith rock outcrops. The steeper slopes are talus coveredas a result of either glacial plucking or the disruptingaction of temperaturechanges in post-glacial time. Glacial lakes formedeitherby dammedup valleysor scoured out basins characterizethe area. For the most part the soils have been derived fromthe weatheringof glacial drift. On the drift overlyingthe High Falls the soil has been mapped as the Lackawanna stonyloam (Jennings,et al 1903). In the depressionson the major ridge, a shallow phase of this type may be recognized. The profile developed is either a true podzol or of the graybrownpodzolic type. The latteris mostwidespread. loam 5-8 in. deep It is a brown to reddish-brown grading into a yellow-brownto Indian red subsoil. The parent materialexposed at a depth of about 2 ft. along the road cut is boulderyclay. It consists of a mottleddark-brownto light-graymatrix containing many bouldersand rock fragments. On the Shawangunkformationrock outcropsare common. Here wheresoil has formedat all it is by the slow process of the accumulationof organic mnaterial and weatheringof the parentmaterials. The park has an annual mean temperatureof 49.5 F. (Sussex, 22-yr.record) with extremesranging from -20? to +1000F. The frost free season extendsfromMay 7th to October1st (CulversLake). The annual mean rainfallis 45.94 in. (CulversLake) withmostof it occurringduringthe summer. Much of the precipitationin winteris in the formof snow with occasional sleet and ice storms. The annual snow coveris 54.5 in. (Culvers Lake) or 10% of the total precipitation. Snow to a depth of 20 in. in one stormis not unusual. The wind directionis southwestduring July and August but from the northwestduring the rest of the year (U. S. Dept. Agr. 1926). The park lies in the oak-chestnutforest region whichextendssouthwardalong the Ridge-Valleyand Blue Ridge Provinces (Blodgett 1910; Braun 1950). To the east on the Piedmontof North Carolina,this forest gives way to oak-hickory (Oosting 1942) whichalso occursnorthwardon the Piedmontof New on Jersey(Bard 1952). To the northand northwest the AlleghenyPlateau, chestnut-oakis replaced by pine-northernhardwoods as dethe hemlock-white scribedby Nichols and others (Nichols 1913a, 1935; Lutz 1928, 1930; Bray 1930; Hotchkiss1932; Hough
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Fim. 2. View north from top of High Point Moniuent in autumn. On ridge at right, oak conununity(lemarked by green areas of pitch pine. Contiguous with pitch pine-scrubb it is chestnut oak communityrepresented by orange-yellow coloration. This community also predominates on the adjacent slopes. In center at base of slope, Cedar Bog with lark green coniferous element (hemlock, black spruce, southern zwhitecedar) and(linterspersed reddish-yellowdeci(luous element (red mal)le, Yellow birch, black gum). Reddish tinge, at base of slope and especially beyond the bog, indivates the presence of other oak species. Grayish-whiteareas on ridge top within pitch pine-scrub oak communityare conglomerate rock outcrops surroundledby scarlet eolored ericaceous shrubs. Dark areas beyond bog are cloud shadows. (This illustration published with funds other than those of Ecological Monographs.)
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April, 1953
PAVST AND PRESENT
VEGETATION
OF
Hioii
POINT
STATE
1PARK
129
FIG. 3. View south from top of High Point Monument. Shawangunk conglomerateforms major ridge which continues southwestward. Less rugged topographyof High Falls formationto right of major ridge. Pitch pineserub oak communityin foregroundleft of road (Stand 1). Note burned-overarea left of center, indieated by carpet-likearea of low shrub vegetation. Lake Marcia at right. Fields are on Martinsburgshale at left. Lake Rutherfordand Saw Mill Pond in left and right background respectively.
E. T. Moul 1936; Gordon 1940; Egler 1940; Hough & Forbes (Picea); W. E. Manning (Car!la); 1943). Bray (1930) studyingthe vegetationof New (Mosses); J. W. Thomson (Lichens); 11. A. Wahl York State, pointsout that the State lies in a tran- (Cares); and F. W. Woods (11re). sition zone whereoak, chestnut,and hickoryoverlap METHODS with the northernhardwoods,the birch,beech,hemlock, and sugar maple. Similar patterns in more The fieldwork was conducte(lduringthe summers detail have been observedby Raup (1938) in the of 1950 and 1951. A systematicreconnaissance of the rugged Black Rock Forest of New York and by park was undertakenby followinga series of comJennings (1939) in the mountainousRidge-Valley pass lines selectedon the basis of a study of aerial Province of Pennsylvania. Within the High Point photographs and topographic maps. Then long region, as early as 1884, Britton noted a striking transectswere establishedthroughsections of the similaritybetweenthe pitch pine-scruboak on the park, cuttingacross successivecomm iiunities. Along Kittatinnyridge and the Pine Barrens of southern these transectstree cover was mleasuredby the line New Jersey. He also observed the presence of a intercept method (Bauer 1943, Buell & Cantlon peculiar floracharacteristic in part of miorenorthern 1950). This served to show the relationshipof the regions,or in part, the coastal plain of New Jersey. conununitiesto the topographyand to each other. Around1900 the geologicaland soil surveysreported There were five such transects (A, B, C, D, E). a mnixed coniferousdeciduous forest (Vermeule, et Next,individualstands were sampled (Stands 1-19). al 1900). Jennings and his co-workersfound a Locationsof transectsand stands are shownin Fig. forest predominantly of oak-chestnut(Jennings,et 1. Again the line interceptmethod was used to of determinecover in both the tree and shrub layers. al 1913). Moore (1939) reportsa forestprimarily in whichoak is dominant. Recent In each community mixeedhardwvoods a line of 100 m. was used except comparativestudieson the life formsin the pine-oak where only a single stand of a type was sampled. community comparedwiththosein the Pine Barrens In the latter case, the size of the sample was inof New Jersey, indicate a greater proportionof creased dependingupon the uniformity of the commihemiciryptophytes and cryptophytes on the ridgethan munity. For instance,in the mixed oak-hardwoods in the Pine Barrens,whichis suggestiveof the more communitya transect of only 160 m. was used. rigorous climate of the ridge (Archard 1953, in Diametersof all trees whichoccurredin at continupress). ous 4 m. strip along the transectline were measured. I wish to express my sincere appreciation an(l Saplings (includingindividualsof tree species I ft. thanksto Dr. and Mrs. Murray F. Buell for their high or over but less than 1 in. d.b.b.) Adelecounted continuous encouragement,sincere interest, and on a continuous2 n. strip along the transectline. helpful criticismsin the preparationof this manu- Herb and moss cover was estimatedoln 1/ x 2 in. script. And also to Mr. Howvell0. Archardfor his quadrats spaced at 5 m. intervalsalong the trisect assistance duringthe fieldwork. I am indebtedto line. Tree seedlingswere countediil the ?i, x 2 iim. the followingfor their identification or verification quadrats. of species of the followinggroups: Drs. G. M. Day Specimensof all species found were collectedand
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130
WILLAM
A.
have been depositedin the ChryslerHerbariumof Rutgers University. Nomenclaturefor the vascular plantsis accordingto Gray'sManual (8th ed. Fernald 1950); mossesaccordingto Grout (1940). studSoil samples were taken in the communities ied. The pH of the samples was determinedwith the Beckman pH meter after the soil had been broughtinto the laboratory. The dry soils were soaked 24 hours before testing using a soil-water ratio of 1:2.5. Peat samples for pollen analysis were obtained fromCedar and Pine bogs. The sampleswere taken at 1 ft. intervalswith the Davis-typepeat sampler. In the laboratorythe pollen was separated by the KOH method(Sears 1930). Counts of 150 grains were made except in the lowerclay layerswhere,because of a paucityof pollen,a minimumof 50 grains was counted. Such low pollen samples come from the 33 to 38 and 40 ft. levels of Pine Bog and the 20 ft. level in Cedar Bog. Percentagesare based on total tree pollen.
NIERING
Ecologicl 2Monographs
the ledge near the monumentwheretheyoccur with Potentilla tridentata(Fig. 4). These three,Bryum, Ceratodon,and Potentillaalso occur in an admixture of soil and conglomeratefragmentssurroundingthe outcrops. Othermosseswhichcommonlyformtufts and mats in the crevicesor depressionsare Polytrich&um juniperinum,P. commune,Leucobryuma glazecum,Dicranumecopariumand D. fulvum. Cladonia, especiallyC. caroliniana(Schwein.) Tuck. may occur with any one or combinationsof these species and eventuallyform expanding mats which spread out over the rock surface. Similar mats formfromthe edge and advanceout overtherock. The thicknessof the moss-lichenmats increases as the lower portions of the mosses and lichensdie and add humus to the organic layer and as dust and dirt sift down into them.
RESULTS XERARCHSUCCESSION The earlierstages of primarysuccessionare characterizedby the coveringof exposed rock surfaces with vegetationand its accompanyingsoil accumulation. The pioneer invaderson the bare rocks are usually crustose and foliose lichens. The formamats occursin the depressionsor tion of moss-lichen from the peripheryof the rock outcrops. As soil accumulateswithin these mats, grasses and sedges FIG. 4. Xerarch succession on Shawlangunkconglomare able to becomeestablished. With furtheraccumu- erate at High Point Monument. Rinodina ori(oa gives lation ericaceousshrubs and trees of the surround- conglomerate gray tone. Umbilicarisceae form black ing forestinvade and eventuallycover the exposed scaly patches at right. In crevices Potentillatriidentata is flowering. Note Pyrua rnelanocarpaat left. rock surfaces. This succession is well shown on the exposed outcrops of the ridge top, contiguous With the increasein thicknessof the mat,growing, with the pitch pine-scruboak community(Fg. 2). conditions improve,especially moistureholding caGrowingon the dryexposedrocksurfacesof theridge top, crustoselichensmustwithstandextremeperiods pacity,and herbscan growon themat. Deschanipsia of drought. Most conspicuousof themis Rinodina flexuiosais among the firstand along with it may Anoreina (Ach.) Mass. whichgrowsso closelyappressed be Danthonia spicata, Panicum depauperatuim, and Carex dropogon scoparius, pensylvanica. Iit and be a it to of part to the rock that it appears gives the conglomeratea blackish-grayappearance creviceson the moreexposed sites,Carex, Dantlhonia. (Fig. 4). Foliose formssuch as Crocynianeglecta and Panicum frequentlyoccur with the mosses an(I (Ach.) Hue. and Parmelia conspersa (Ehrh.) Ach. lichens. During this open herbaceous stage pine var. isidata (Anzi.) Stizenb. also grow on the bare seedlingsbecome established. Finally, with further rock surfaces where they frequentlycover and de- growth of the mats, low shrubs invade either on stroy the crustoseforms. Crocynia appears as ir- the peripheryor in the centerof the mat. In order regular gray patches and Parmelia forms circular of decreasing importance on the mats the! are greensh gray patches. In additiontwo membersof Vacciniuimangustifolium,Gaylussacia baccata, PyActinogyramuhlenbergii(Ach.) rus melanocarpa, Vaccinium vacillanes,Kalmniaanthe Uimbilicariaceae, Schol. and Lasallia pensylvanica (Hoffm.) Llano gustifolia, Amelanchierstoloniferaand Vaccinian formiblack scaly masseson the rock surfaces. These staniineum. Where soil accumulatesin the deeper lichensdo not developmats upon whichhigherplants fissuresof the bed rock,shrubsmay invade directly. can grow. The soil buildingpioneerswhichstartin Here Pyrus or one or more of the above shrubs is the crevicesand depressionsor fromthe periphery frequentlyfound. Scrub oak (Quercus ilicifolia) of theoutcropshere,as in NorthCarolina (Oosting& becomes establishedon the mat only after it has Anderson1937), do not appear dependentupon the reachedconsiderablethickness. Herbaceouscoverbepresence of crustose lichens.Bryum capillare and neaththe shrubsis similarto thatfoundin the pitch Ceratodonpurpurens are theearliestformsto appear pine-scruboak community. In contrastto the exposed ridge top is the northin the crevices. They are particularlyimportanton
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April, 1953
PAST AND PRESENT
VEGETATION
OF HIGH
POINT
STATE
131
PARK
occasionally Rhododendron maximum. The trees whicheventuallybecomeestablishedgrow withtheir roots spreading throughthe moss mats. However, as the roots increasein size and portionsof the mat are eroded away, the roots becomeexposed. These rootsof the largertreesfrequentlygrowbetweenthe boulders. The trees,thus poorlyanchored,frequently lean and oftenexhibitinferiorgrowthform. On most rockyareas this successionlong ago reacheda forestedstage resultingin many rocky woodlands in the park. 2'~~~~~~~,
HYDRARCH SUCCESSION
Two patternsof hydrarchsuccessionare evident in the park. In the artificialimpoundments, filling is occuringprimarilyby sedimentationand by the invasion of emergentaquatic vegetation. In contrast,lakes of glacial origin are being filledby the encroachmentof a floatingmat upon which vegetation thrivesand which deposits organic material from the under side. However, in both of these patterns,submergedand floatingaquatics are presA~~~~~~~ ent in the deeper water. In the two instances,the of emergentsor by mat forfillingby encroachment mation,different types of forestcommunities result. In the formerthe forestis composedprimarilyof broad-leavedtrees (swamp forest), in the latter of conifersand scatteredbroad-leavedspecies (bog forest) (Fig. 1). Both of these successionalpatternsare found in the park area. The earlier stages of the succession may be observedin the five lakes and two beaver FIG. 5. Northwest facing talus slope above Cedar Bog. Note sweet birch and white birch. Rhododendron dams. Three of them,Steeny Kill Lake (No. 20), Saw Mill Pond (No. 21), and an unnamed pond is doi'uinantshrub. (No. 22), are artificialimpoundments(Fig. 1). In Lake Marcia (No. 23), and Lost Lake (No. contrast west talus slope (Fig. 5). There muchof the slope 24), are of glacial origin. Sussex reservoir(Lake oak chestnut is partiallyshadedby (Quercus prinus), Rutherford)althoughsurroundedby park property sweetbirch(Betnla lenta), whitebirch (Betula papywas not studied. strobus), striped maple rifera), white pine (Piuns (Acer pensylvanicuin), mountain maple (Acer spicaSuch tu1ni) and mountain ash (Pyrns americana).
boulderson slopes are strewnwithlarge conglomerate which the earlier lichen stages are similar to those on the ridgeexceptfor an abundanceof Umbilicariaceae. Amongthese Lasallia papulosa (Ach.) Llano is doninant, forning large grayish-blackscale-like growths. In additionUinbilicaria mammulata (Ach.) Llano and Actinogyra muhlenbergii are present. Where the boulders are shaded, these species increase in cover so that littleof the rock is exposed. On and betweenthe rocks wheredebris accumulates, mats of Cladonia rangiferina (L.) Web. and C. uncialis (L.) Web. are commonand frequentlyassociate(l with cushions of Polytrichum ohioense, P. COMnI mane, Leucobrya(m glazicum, and Dicranam scoparbinmwhichoccur in pure or mixed aggregations. leaf In thesemoss-lichen cushions,or wheresufficient in and oclitter accumulates,Poly podlia n ritrginianu casionallyDryopteris mnarginalisand A ralia nudica alis are present. Birch seedlings are coinnionon the cushions, as are also such shrubs as Gaylussacia bacc{a, Vacciinna, uagnstifolinni, Kalinia latifolia. and
IIYDRARCIH
SUCCESSIO.N
WITHOUT
A FLOATING
ALAT
Aquatic zonationis whell illustratedat the southern
end of Steeny Kill Lake (Fig. 6). The area occupied by the lake wvasformerlya swamp forest. The trees
werecut and a dani builtat the northern end. Stumps project above the water in the southernend of the lake; elsewherethe deeper waterhas submergedthem.
Callitriche heterophylla and species of Potamogeton are the predominant submerged forms. On the exposed stumps grow Leersia oryzoides and Spiraea tomentosa. In the shallower water, is an emergent zone dominated by Sparganiumr americanum which grows 1-2 ft. in height (Fig. 6). Its associates are Iris versicolor, Leersia oryzoides and several species of Scirpus. On hummocks in this zone Spiraea occurs with Hypericitm vijighiicuim,H. mutildit amld Galiunm tinctoriunr. Further back Juncus effusIus and Scirpus cyperinus becoimme the principal species. Between the hummocks of Carices and Osinunda ciazmanmonmea, Sphagnum forms a more or less continuous layer with scattered plamntsof Drosera rotmimdifolia, Dryopteris thelypteris. Clhrysospleuiem ancri(CliUM0, and Sciilpis (irochiiht s. Here in(d there
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132
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Ecological Monographs Vol. 23, No. 2
way to Alnus toward the upland. In other places along the marginRhododendronviscosum,Rex verticillata,Vacciniumcorymbosum, Viburnumrecognitum, Alnus serrulata,Alnus rugosa and Cephalanthus occidentalisforma dense border. In the open water,herbaceousaquatics are relativelysparse. In the shallow water near the shore one findsa few scatteredpatches of Ludwigia palustris. In the southernpart of the park, beavershave built a dam, forminga pond in a low swampy area, and killing many swamp hardwoodspecies. Along the pond margin Typha latifolia grows in sporadic clumps. On the surfaceof the waterone frequently FIG. 6. Hydrarchsuccessionat southernend of Steeny findsa dense coveringof Lemna minor. In the cenKill Lake. Note concentric zones beginning with tral part of the park, beavers have recentlyflooded Sparganiuimamericanumat edge of water and followed a low swampyarea and the surroundingoak forest. by Juncus-Scirpusstage, shrub stage, and finallyswamp Sonic of the trees are dead and others are dying hardwoods communityin left background. eitherdue to floodingor the activitiesof the beavers girdlingthe bases of the trees. Aquatic vegetation are pure stands of Typha latifolia in the Juncus- is sparse due to the recencyof the disturbance;howScirpus zone, especially near the shrub zone which ever, near the shore Lemna minorfrequentlycovers is further back. the surfaceof the water. A shrub zone occurs next to the sedge zone and in Anotherarea, at one time inundatedby the activit Alnus rugosa, Ilex verticillata, Nemopanthus mu- ities of beavers and since abandoned, has been cronata, and Vaccin iUnz corymbosum are predomi- drained and today Big Flat Brook flows through nant. Less important shrubs are Rhododendron it. The area showsevidenceof havingbeen pastured maximum and Rhus vernix. The shrub layer is from before flooding. After its use as a pasture ceased 3 to 10 ft. in height. The wet, hummocky ground graybirch,red maple and hawthorn(Crataegusspp.) is covered with Sphagnum and herbs: Polygonum invaded. The floodingproducedby beaver activities sagittatum, 0smunuda cinnamiomteu, Dryopteris the- killed the birchand hawthorn. Since the withdrawal lylpterisand various species of Carex. Young trees of water, a dense willow thicketnow grows along of red maple (Acer rubrum) and gray birch (Betula much of the stream. Rushes, sedges, and grasses ))opulifolia) are especiallycommonin some parts of are also importantalong the streamand on the exthe shrub zone. posed lake bed along withspecies of Eleocharis and At other places along the lake shore where the other herbs. In soire areas the herbs give way to bottomgradientis steeper,the zonation is less con- shrubs-Viburnumdentatum,V. lentago,Lyonia lispicuous and somewhattelescoped. In places Ce- gustrina,and Alnus rugosa. phalanthus occidentalis and Scirpus validus play an All of these earlier stages in hydrarchsuccession importantpart in the succession,but are scatteredin appear to be leading toward a swamp hardwoods their distribution. community.Stands representingthis community ocIn contrast,probably because of the rocky sub- cur in poorly drained areas, chieflyalong the tribustratum,aquatic vegetationis poorly representedin taries of Big Flat Brook. Here two stands (14, 15) Saw Mill Pond and Lake Marcia. In the former were studiedin detail (Fig. 1). Sparganiuim americanum and Brasenia schreberi are The forestflooris wet and boggy. In one stand sporadic along the shallow lake shores. At Lake (15) a mixtureof woody,sedge,and sphagnumpeat Marcia, those plants which occur are restrictedpri- extends to a depth of 3 ft. This is underlain by marilyto shallowinletswheremuckysoil has accumu- yellow-brownsedge peat which gives way to gray lated. Here one may findpatches of Juncus effusus, clay at a depthof 9 ft. J. acutmintatus,Eriocaulon septangulare, Ludwigia The tree canopy is composedof two moreor less 1walustrisand species of Eleocharis. Other species distinct layers, below which is a sporadic shrub that have been found in Lake Marcia but contribute layer and fairly continuousherb and moss layers. littleto the vegetationare Juncus canadensis, Scirpus In the treelayer,red maple and yellowbirch (Betula ralidus, and Eleoclharis palustris (Renlund 1951). lutea) are the dominantspecies, contributing up to Along the rocky shores a dense shrub thickethas 70% of the total tree cover in the two stands. In developed with Vacciniunm corymbosum, Rhododen- one stand, yellow birch is most important;whereas dron riscosum, Myrica gale. Celphalanthus occiden- in the other,red maple is predominant. Associates talis, Clethra (ainifolia and A/lmvrugosa as the princi- are black ash (Fraxinus nigra), whiteash (Fraxinus pal species. americana), basswood (Tilia americana), and tulip In the southernsectorof the park, a small shallow tree (Liriodendron tulipifera) (Fig. 6, 13). The untnained pond is surroundedby a dense shrubzone. tree canopy covers 94% of the total area. Most of At the easternend on the exposed muddysubstrate the trees are 4-9.9 in. d.b.h. although larger ones a willow thicket(Salix serice() is dominant,giving rttaining16-18 in. d.b.h. are represented. In gen-
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April, 1953
PAST AND PRESENT
VEGETATION
OF HIGH
POINT
STATE
PARK
133
eral, the trees are shallowly rooted and somewhat elevatedabove the frequentlywater-filled depressions on the forestfloor. The average heightof the forest Ais 65 ft. In the understoryhornbeam (Carpinus caroliniana) is most important,with hop hornbeam of the domi(Ostrya i-rginizana) and transgressives nants. Yellow birch,red maple, whiteash, and hop hornbcaimi are all reproducingthemselves. Yellow birch scedlings are il!ost abundant followedbv red maple and whiteash. In the sparse shrub layer, covering16% of the total area, there is considerablevariation. In one FIG. 7. Lost Lake in Pine Bog surroundedby a floatstand (14) HIamamelis virginiana and Vaccinium corymbosum contribute71% of the total shrubcover. ing mat on which Chameedaphne calyoulata is the dominant species. Tamarack and black spruce in foreground. In the other(15) Viburnum dentatum and Viburnum Peat samples were taken on the opposite side of the lake lentago contribute57%. Shrubs present in both where the person is standing. stands are listedin figure14. The herb layer is relativelycontinuous,covering tamarack (Larix laricina) have been seedcd in from about 70% of the area. The three most important the surroundingtree and shrub zone. The tamarack herbs in each stand are Osmunda cinnamomea with is restricted primarilyto the open mat; the black a 33% contributionto the cover, Carex stricta with spruce is not only on the mat but throughoutthe 29%, and Symplocarpus foetidus with 5%. Carex greaterportionof the moremiaturebog. Along with and Osmiundaform characteristichummockson the black spruce,red maple,black gum,yellowbirchand forest floor with Sphagnum occupying the wetter sporadic white pine and hemlockforma very open depressionsbetweenthem. Otherherbsare listed in forestrisingabove a dense shrub layer. The shrubs figure15. are mostly Vaccinium corymbosum, Rhododendron Bryophytesoccupy31% of thearea. Sphagnumis viscosum, Alnus rugosa and species of Ilex. Rhodomost important, contributingabout 21% cover. dendron maximum formsa borderalong the edge of Others,in order of decreasing importance,include the bog. Thuidium delicatulum, Mnium punctatum var. elaA later stage in bog succession is illustratedby tumr,Mnium cinclidioides, Tetrapis pellucida, DicraCedar Bog (locally called Cedar Swamp), which is num scoparium, Bazzania trilobata and Trichocolea locatedat the northernpark boundaryapproximately tomentella. They are usually found on the elevated 1500 ft. above sea level (Fig. 1 Stand 19). It is tree roots and sedge hummocks. probable that this was a lake that filledin the same Near Lake Rutherfordthe swamp hardwoodscom- way as Pine Bog. However,the encroachment on the munityoccurswithred maple,black gum (Nyssa syl- lake is completeand no open water remains. The vatica), and sporadic whitepine whichtowersabove area is now completelycoveredby a forestof conifthe deciduousspecies. Vaccinium corymbosum,Rho- erous and deciduousspecies. Althoughit is mostlya dodendron viscosum, Alnus rugosa, and A. serrulata rathcr closed forest,the southwestportion is like are the dominantshrubsthereand gray birchforms muchof Pine Bog withred maple,yellowbirch,black a conspicuousborder. Frequentlyhemlock (Tsuga gum and occasionallywhite pine and black spruce, and black canadensis) gum are mixed in with the scatteredthroughouta shrubbymatrix. other broad-leavedspecies as at the northernand Where the bog forestis most mature (Stand 19) southernends of Steeny Kill Lake. In otherpoorly it consists of distincttree, shrub and moss layers. drained areas, swamp-whiteoak (Quercus bicolor) There are very few herbs. In the tree canopy,hemis also present. lock and red maple are dominant,contributing 55% HYDRARCH SUCCESSION WITH A FLOATING MAT of the total tree cover. Other trees contributing The second patternof hydrarchsuccessionis asso- cover are southernwhitecedar (Chamaecyparis thyciated withthe formationof a floatingmat. This is oides) with 14% cover, yellow birch 12%, black best illustratedby Pine Bog (locally referredto as spruce 10% and black gum 9% (Fig. 8, 21). Cover Pine Swamp) located on the southwestern boundary in the tree layer is 89% withonly 11% represented of the park, 1200 ft. above sea level (Fig. 1, No. 25). by openingsin the canopy. The largerhemlocksand It was formedas a result of glacial driftblocking cedars are approximately, 90 and 140 years of age the drainage of a mountainvalley (Waksman, et al respectively. The tops of manycedars and hemlocks 1943). In the southernpart of the bog, an open are brokenor dying back as a result of wind and body of water (Lost Lake) is surroundedby a float- ice storms. Others,especiallyhemlock,are branched ing sedge and sphagnummat (Fig. 7). On the mat, on the side oppositethe prevailingwinds. Windfalls Chamaedaphlne calyculata is the dominantshruband are occasionallyfound with the shallow root systems along with it are Andromeda glaucophylla, Kalmia of the fallen trees tipped up on edge. Logs in polifolia, K. angustifolia, and Vaccinium oxycoccos. various stages of decompositionfurnishan excellent Here smalltreesof black spruce (Picea mariana) and substrateforestablishment of seedlings. Dead stand-
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1:34
WILLIAAM WILLIAM
A.A.NIERING
Ecological Monographs 2
~~~~~Vol. 23- No-
palustris, Sarracenia purpurea, Drosera rotundifolia, Scirpius cyperinus, Eriophorum sp., Juncus effusus and species of Carex and Eleocharis. DEPOSITS
FIG. 8. Cedar Bog. oi(IeS most conspicuous.
Crownsof Chamaecypari8 thy-
Road crosses bog at this point. Youing ced irs Ire common along the road.
OF PINE
BOG AND CEDAR
BOG
Samples of the sedimentsnow fillingthe lake basins of Pine and Cedar Bogs were obtained to a depthof 40 ft. and 30 ft. respectively.In the lowest levels, the deposits consistof 8-10 ft. of gray clay. These are overlainby gyttja whichis formedin the open lake waters as a result of benthicfauna reworkingplanktogenicdetritus(Lindeman1941). The gyttja varies from fine detritusooze in the lower levels to coarse detritusooze in the upper levels (Deevey 1939). Its depth ranges from10-15 ft. in the two bogs. The upper layersof sedge and sphagnum peat were formedby dead materialof sphagnum,sedges,and associated plants falling fromthe undersideof the mat. As this is continuouslybuilt up by fresh deposits,it compressesthe underlying gyttja. In Pine Bog 17 ft. of sedge-sphagnum peat has accumulated. In Cedar Bog thereare 4 ft. of sedge peat containingfragmentsof wood, overlain by 6 ft. of woody peat.
ing trees,especiallycedars,are scatteredthroughout; one cedar measured25" d.b.h. Some cedars up to 30" in diameterhave been cut. The trees differin theirability to completetheir life cycle. Hemlockand red maple are mostsuccessful. Cedar seedlingsare sparse except in the openings. Those that becomeestablishedin dense shade usuallydie when3-4 in. in height. Cedar is destined ABANDONED FIELDS to be of less importancein the futureas a resultof Most of the formerfarm land is now abandoned competitionfrom the nmoreshade toleranthemlock to the invasion of native vegetation,and various and red maple. Only locally is it increasingin im- successionalstages are evidentleading to an oak or portance. For example,along a road recentlycon- oak-hickoryforestcommunity(Fig. 1). Along the structedacross the bog, whitecedar shows vigorous lowersoutheastescarpment, on the Martinsburg shale, reproductionwith an abundance of seedlings and abandoned fieldsare more or less continuous. Erosaplings and largertrees3-4 in. d.b.h. Black spruce sion there has removedmuch of the glacial drift, reproductionis also poorly represented,it, like the exposing the shale so that the vegetationthere is cedar being moreimportantin the less matureparts directlyinfluencedby the underlyinggeological forof the bog. Reproductionof yellowbirchand black mation. On the High Falls and Shawangunkforgumuindicatesthat they will maintaintheir present niations,throughoutthe rest of the park, the old status in the community. fieldsare morescattered. Wherethereare old fields, The shrublayer is 93% complete,therebeing only glacial driftcovers the bed rock. Agriculturaluse 7% of the area unoccupied. Of this shrub cover, of the land has ceased, except for several fields Rhododendron maximunlm, whichoccurs primarilyin seeded withtimothyand clovera few years ago and deepershade,formsa denseundergrowth contributing still iuowedeach summer. 64%. Its seedlingsare abundanton the moss-covered PERENNIAL STAGE HERBACEOUS hummocksand root systemis. The remaining36% There are no fieldsjust abandoned,but on those of the shrub cover is contributedby Ilex laevigata, I. verticillata, Nemopanthus mucronata, Vacciniurn with good drainage in which cultivationhas most and Alnis serrulata which occur pri- recentlyceased, Poa cornpressa, Daucus carota, Solicorymnbosurn niarilyin the openings. In the southwestportionof dago graminifolia, and Hypericurm perforatur are the bog whereit is more open Rhododendron maxi- the dominants (Fig. 9). Next in importanceare mum frequentlydecreases in importance. Here in Hieraciurm pratense, H. aurantiacurm,Trifoliurmpraadditionto thoseshrubspreviouslymentionedRhodo- tense, T. repens, Linaria vulgaris, Agrostis alba, dendron viscosum, Alnus rugosa and Lindera benzoin Phleirm pratense, and about 34 otherherbaceousspecies scatteredthroughthe vegetation. Andropogon forman almostcontinuousshrublayer. In the mature forest,herbs are widely scattered. scoparius is not very importantin the succession except in fields whose soils are derived from the Osmnunda cinnamnomeacontributes5% cover, Calla palEstris 3%, and Symplocarpus foetidus 1%. These shale. Fields in this early stage of successionare bordered by hedgerowsof gray birch, are mostimportantin the openings. Of the remain- cotmmonly ing 13 herbs each contributesless than 1% cover. tremblingaspen (Populuis tremrldoides),black cherry Beneath the shrubbymattrix in the more open por- (Pr.nus serotina), scarlet oak (Quercus coccinea), tions of the bog, tussocksof Osmunda cinnamomea and red maple (Fig. 9). From this border the are characteristic.Alongthe trees spread out into the fieldand one findsyoung coveredwithSphagnunm road across the bog, Sphagnumsupportsa distinctive plallts among the herbsnear the edge of the field. In the southernpart of the park on the poorly herbaceous flora including species such as Calla
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April, 1953
PAST AND PRESENT
VEGETATION
OF HIGH
POINT
STATE
PARK
135
(Fig. 10). Some of the saine shrubs found in the thicketpersist but there is a change in the herb composition. The principal ones are Lycopodium
complanatum, Pteridium aquilinum, Lysimachia qua(irifolia, and Medeola virginiana. The resultingforest
is dominatedby eitherred oak, chestnutoak, white oak and hickoryor by whiteoak and hickoryalone.
FIG. 9. Perennial herbaceous stage. Daucus carota and Soli(lago graminifolia are dominant herbs. Gray birch aend trembling aspen at left and oaks at right borderalong hedgerow. formn
drainedPapakating silt loam,Potentilla simplex, Iris versicolor and species of Carex, Juncus, Solidago, and Agrostis are dominantin the fieldsthat have not been plowed for some time. Other species present are Linaria vulgaris, Hypericum perforatum, Cirsium arvense and Phleum pratense. Woody species of foreststage. Graybirchand FIG. 10. Approaching Spiraea and Clethra are invadingthiscommunity. pitehpine still present. Largertreesare oaks. THICKET
STAGE
Gray birchspreads out over most fieldsrelatively Other trees commonin the young forestare biacL fast (Fig. 9). Along with it appear the shrubs- oak, red maple,and scarletoak. Rhu1s typhina, R. glabra, Quercus ilicifolia (scrub
oak), RClbus spp. and various heaths. The herbsof this thicketstage are primarilyAgrostis alba, Poten-
tilla sinmplex,Solidago graminifolia, Hypericum perLinaria vulgaris, Solidago rugosa and Poa foratutmn, cornmpressa.As the successioncontinuesconsiderable
variationis evident.Graybirchmayforma relatively uniformand continuous,treelayergrowingin clumps and rangingfrom 1-4 in. d. b. h. With it may be found scatteredblack cherry,red maple, and white ash as well as saplings of red oak (Quercus rubra), white oak (Quercus alba), red maple, and scarlet oak. Species of Rubus commonly forma densetangle 3-4 ft. in height. The dominantherbs are Solidago rugJOS(l and Potentilla simplex. On the otherhand,gray birchand red maple may in whichcase theshrublayermayconbe co-dominant sist of Quercus ilicifolia, Vaccinium angupstifolium,
PITCH PINE-OAK COM.MUNITIES
The pitch pine-oak communitiesoccur on the Shawangunk conglomerateas a depauperate pinescruboak community (Fig. 11) and on thedriftoverlyingthe High Falls sandstoneas a pitch pine-arboreseentoak community.Four standsof the pine-oak conmiunitywere studied,two (Stands 1, 2) on the and two (Stands 3, 4) on Shawangunkconglomerate the High Falls sandstone. The formerwill be referredto as the pine-scruboak and the latteras the pine-oak.
V. stamineum, Viburnum recognititm, Hamamelis virginiana and species of Rubus. In such stands Lyco podium complanatum and Polytrichum commune
frequentlyform continuousmats over the ground. Where rockoutcropsin the thicket,Polytrichum commune, P. juniperinum and Danthonia spicata cover the area. Sometimeswhite pine and pitch pine FIG. 11. Pitch pine-serub oak community.Note (Pbius rigida) are associated with red maple and crevices of conglomerate. Clhestnutoak coin,graybirch. Saplings of hickory (Carya spp. in- hteatlhsin is contiguouswithpitcl pine at right (Stand 2). 111111uity elu(ling C. glabra and C. ovalis; Carya ovata) red, white,scarlet,and black oak (Quercus velutina) are PLNE-SCRUB OAK aLso common. On the Martinsburgshale red cedar On the ridge where the pine-scrub oak com(Juniperus virginiama) is occasionallyassociatedwith mnunity occurs,thesoil is verythinand rockoutcrops gray birch and red maple but does not occur in the are common(Fig. 11). Often mnats consistingpriother areas.
FOREST
STAGE
Eventuallya forestof oak or of one dominatedby oak
and
hickory
gains
ascendancy
over
the birch
mnarilyof organic material bound together with roots cover the rock surfaces. The pH is approximately 4.
The tree and herbaceouslayers are open, in contrastto a shrublayer that furnishespracticallycom-
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136
WILLIAM
A.
plete cover (Fig. 13, 14, 15). The pitch pines, the dominanttrees contributing76% of the total tree cover, are ratherwidely spaced. Species associated with the pine in order of decreasingimportanceare red maple, gray birch,juneberry (Amelanchier arborea, A. laevis), and sweet birch. The tree layer averages 18-20 ft. in height with the larger pines 8-10 in. d. b. h. and approximately70 years of age. The pines typicallyhave brokentops or tops with gnarledor twistedbranches. The open aspect of the woods is evidencedby the 54% unoccupiedspace in the tree layer (Fig. 13). Pine seedlingsand saplings are oceasionallyfound soil in the rock crevicesor moss mats wheresufficient is present. Seedlings of red maple are frequently found. Saplings of sassafras (Sassafras albidum), juneberry,red maple, and sweet birch are scattered throughoutthe stands. Although a chestnutoak communityalways bordersit, chestnutoak saplings of the are rarelyfoundin the pitch pine community ridge top. The shrubs are conspicuouslytwo-layered,covering 92% of the area. The taller shrubs are scrub oaks whichoftenforma dense thicket3-10 ft. high. The lower layer is dominatedby heaths, of which Vacciniurn angustifolium and Gaylussacia baccata, typicallytwo feet or less in height,are most important contributing57% cover. Low shrubs of less importanceare Pyrus melanocarpa, Vaccinium vacillans, Kalmia
angustifolia, Amelanchier stolonifera
and Comptonia peregrina, Diervilla lonicera, Prunus pensylvanica, P. pumila, Rhus copallina, and Pyrus americana.
The herb layer, in contrast,covers about 20% of the ground. Of the 21 speciesof herbsMaianthernum canadense, Pteridium aquilinum, and Aralia nudicaulis togethercontributeapproximately20% cover in this layer. All the others contributea total of less than 1%. The moss layer is sparse covering less than 1% of the area. Evidence of browsing by deer is conspicuous. Plants most frequentlybrowsedinclude pitch pine, sassafras,juneberry,mountainash saplings and certain herbaceousspecies. Evidence of firein the recent past is indicatedby the firescars on the base of the tree trunks. PINE-OAK
The pine-oak occurs on some of the hilltops of the High Falls formation. The soil is a podzol, the A horizonash-grayin color underlainby a yellowbrownB horizon. The pH of the soil is about 4.5. The treecanopy frequentlyconsistsof two moreor less distinctlayers. The uppermostlayeris dominated by occasionalpitchpine extending10-15 ft. above the lower oak canopy. The pitch pine contributesapproximately24% cover. The deciduouslower layer of red maple, scarlet (Quercus coccinea), white,and chestnutoaks representsabout 65% of the total tree cover (Fig. 13). Trees of minor importanceare black gum (Nyssa sylvatica), black oak (Quercus relutina) and sassafras. Thereis about 20% unoccu-
NIERING
E Vo1lg.ca M23 Nograph
pied space in the tree layer comparedto about 50% in the pine-scruboak communityof the ridge top (Fig. 13). The larger pines are 17-19 in. d. b. h. and about 140 ycars of age. Most of them are over 10 in. d. b. h. The hardwoodsare most abundant in the small size classes. Pine snags and rotting pine logs occasionallyare present. White oak, scarlet oak, chestnutoak, and red maple are reproducing, and chestnutsproutsare present. Pine seedlingsare rare and saplings are completelyabsent. In the shrub layer, Gaylussacia baccata is dominant, contributingabout 60% of the cover. Other heathsrepresentedare similarto thosefound on the ridge (Fig. 14). Scrub oak contributesabout 15% of the cover and occurs in clumps, where ample light reaches it. The shrub layer is rather dense, coveringabout 85% of the area. Ten herbs wcre found. They occupy only about 15% of the area (Fig. 15). Pteridium aquilinum has an average of 9% cover. None of the othersfurnish over1% cover. Principal amongthemare Gaultheria procumbens, Aralia nudicaulis, and Medeola virginiana. Mosses occupy less than 1% of the area. CHESTNUT
OAK COMMUNITY
The chestnutoak communityis the characteristic of the park. It formsa matrixin which community all the other coninuniticsare found (Fig. 1). A total of 6 stands was studied. Four (5, 6, 7, 10) of the stands were selectedon the Shawangunkformation; two (5, 10) on the southeastescarpmentbelow the monument;another (6) on the northwestslope above Cedar Bog and the remaining(7) on the ridge above, where it occupies a slight depression. The two stands on the High Falls (8, 9) are located on gentlyundulatingtopographyoverlainby glacial deposits (Fig. 1). On the upper and mid-slopesand manysites along the ridge top of conglomeratewherethe soil is very rocky,the mostxeric aspect of the chestnutoak comHere an organic munityis characteristic(5, 6) layer consistingof 3-6 cm of leaf litter overlies a mor humuslayer whichformsa mat over the rocky substratum. Depressions along the ridge top, filled with soil, constitutea more favorable environment for this community(7). An especiallyfavorablesite for the communityis on the High Falls formation where the soil is relativelydeep overlyingthe bed rock (8, 9). On the lower slopes, although the ground is very rocky,the intersticesare filledwith finersoil resultingin an even more favorable soil mantle than upon the slopes and ridge tops (10). the soil pH is 5.0. In this community Within the chestnutoak community,there is a progressionfrom the most xerophyticaspect (5, 6) (Fig. 12), to the deeper soils, especiallyon the High Falls and depressionson the ridge top (7, 8, 9), to the lower slopes of the conglomeratewhichrepresent the mostmesophyticphase (10) (Fig. 13, 14, 15). On the rocky slopes, the forest consists of two more or less distincttree layers, a relativelycontinuousshrublayer and sparse herband moss layers.
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PAST AND PRESENT
April, 1953
VEGETATION
,-.M -~~~~~~~~~4
FIG. 12. Chestnutoak communityalong road to Cedar Bog. The most xeric aspect characteristic on rocky slopes.
In the tree layer, chestnut oak is by far the most important, contributing over 55% of the total cover. The principal associate is red oak. Black oak and sweet birch are occasionally present. Although the total cover in the tree layer is over 90%, the tree crowns are thin, transmitting a considerable amount of light. MNostof the trees fall into 1-3.9 in. and 4-9.9 in. size classes, but larger specimens are scattered throughout the stand. Two or more trees growing PINK NANis
PIKE
OAK
?113
OKINE
WINS
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OAK
OAK
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OM
K.."N30 gNo"""
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AC9R MINSLVA
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COBSM
CJUNA
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M*AXIMIA
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fofrstom
VACCUUMV
ULMUM MUSM NTS
SYLVTICAW
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FIt. 13. Tree cover and frequency for forest coyof bar represents%w of total coter, munities. L(ngta o c uidthof bar retresents frequencyof occurrenceSon 0 m segmentsalong the transectlines in all stands sampled pine-scrub oak of each comsunity. For example, in the a is dominantcontributings6pac ritig communityPinus of the total tree coaer and representing93% frequency. The (X ) indicates that the species wlas present in the communitybut not occurring oll the transects. Space refers to the portion of the canopy between the crowns of the trees. For examlple;In the pine-scruboak coimmunity,where the gaps ill the canopy are greatest,space is equal to 54%r,of the total area, in contrast with the mixed oak-lhardwoodswhere the canopy is completely closed. Carya spp. include C. glabra and C ovalis.
OF HIGH
POINT
STATE
PARK
137
froma single root systemare common. The larger chestnutoak are 9-14 in. d. b. h. and approximately 95 years of age. The average height of the forestis 50 ft. The understoryconsistsof red maple, sassafras,chestnut sprouts,and young trees of hickoryand oak. All species are reproducingthemselvesalthoughchestnut oak is mostsuccessful. Oak reproductionis primarily of sprout origin. Chestnutsprouts are common. Numerousfirescarredchestnutsnags litterthe forest floor. The shrublayer covers40% of the area. The uniformdistributionof shrubstemsformsa fairlycontinuouslayer. In one stand Gaylussaciabaccata and Vacciniumvacillans are most important,whereas,in the other Kalmia latifolia and Rhododendronnudifiorumpredominate. Othershrubsof importanceinand Kalmis angusticlude Vacciniumanguwstifolium folia. Of the 12 shrubspresent7 are ericads which furnishover 90% of the cover in this layer. The herb layer is extremelysparse, the total cover in this layer being less than 10%. Of the 17 species presentin the herb layer only two,Aralia nudicaulis and the sub-shrubGaultheriaprocumbens,were estimore than 1% cover. Others mated as contributing occasionally present are Carex pensylvanica,Pteri(ium aquilinum and Melampyrumlineare. MIosses contributeapproximately2% cover. The principal species, Polytrichumohioenseand Leucobryumglaucmu frequentlyoccur betweenexposed rocks where the leaf litteris relativelythin or absent. This xeric phase extends onto the ridge tops whereverthe soil is very rocky. Here the trees are smallerwithlow gnarled branches. They frequently occur in clumps of 3-6 from a single root system. ericaceousshrublayIn additionto the characteristic er, clumps of scrub oak occupy the openings in the treecanopy. On the ridge top whereconsiderabledrifthas been depositedin the depressions(7) and over mostof the undulating topography of the High Falls (8, 9) wherethe soil is relativelydeep, oaks such as white, red, and black are associated with the chestnut oak. On thesemorefavorablesites chestnutoak contributes about 43% cover while its associates contribute 24%. The treesfall primarilyinto the 1-3.9 in. and 4-9.9 in. size classes except on the High Falls where the 10 in. or over size class is of equal or greater importance. The largertreeson the ridgeare 8-9 in. d. b. h. and 60-70 years of age while those on the High Falls are 15-16 in. d. b. h. and 110 years old. The average height of the forest is 45 ft. on the ridge and 65 ft. in the High Falls stands. A distinctunderstoryof red maple and.oak transgressivescontributesthe greatestcoverin the understoryalthoughsassafras,hickory,and juneberryare also present. All species appear to be reproducing yet mostof the reproductionis of sprout themselves, origin. Chestnutsproutsare common. Many hickory saplings exhibit inferiorgrowth'due to the dying back of the terminal buds. Many of the larger
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138
WILLIAM
A. NIERING
Ecological Monographs Vol. 23, No. 2
tree canopy contributes97% cover in contrast to 90% on the upper slopes. The larger trees are chieflyoaks ranging from 17-23 in. d. b. h. Red oak exhibitsthe best growthwith straightboles and Lwelldevelopedcrowns. In contrast,the larger hickories frequentlyare somewhatdecayed at the base and thetops are oftenbroken. The averageheightof the forestis 60 ft. as comparedto 50 ft. furtherup the slope. The understoryis similarto the poorer stands exeept for the pronouncedincrease in hickorytransgressives and the presence of floweringdogwood (Cornus florida) and stripedmaple. Hickory,chestnut oak, and red maple are reproducingthemselves, with hickorymost abundant. Saplings of ash and sweet birch are present. Sprouts of sassafras and chestnutare common. Red oak is absentin the seedling class but present in all other size classes. Hickoryand chestnutoak are the only species in all size classes. The shrub layer covers50% of the area. Of the 62% of FIG.. 14. Shrub cover and frequencyfor forest com- 12 shrubspresentCornus rugosa contributes munities. For explanationsee legend for figure13. the cover. Others'of less importanceare Corylus Rhododendron roseumis presentin chestnutoak com- cornuta, Vaccinium vacillans, V. stamineum, Cornus munitybut it was not recordedin the transect data. alternifolia, and species of Rubus. Two vines,Parthenocissus quinquefolia and Vitis aestivalis, contribchestnutoaks are not healthy,and in fact dead trees ute 9% and 4% cover respectively. are occasionally found throughoutthe stand. The herbaceouslayer coversonly 12% of the area. The shrub layer is more or less continuous,but Of the 30 speciesrepresentednot one contributes over considerablevariationoccursamongthe dominantsin 1% cover. Collinsonia canadensis, Solidago caesia, the different stands. The total cover contributedby Desmodium nudiflorum, and Smilacina racemosa the shrubs in approximately56%. On the ridge, contributea total of 3% cover. Otherherbs,in order Vaccinium angustifolium and Gaylussacia baccata of decreasingimportance,include Amphicarpa braccontributeover 80% of the shrub cover, whereas teata, Uvularia perfoliata and Galium lanceolatum. on the High Falls the latter,and Vacciniumvacillans In consideringthe threeaspects of this community, occupy over 75% of the area. Other shrubs repre- chestnutoak is the characteristicspecies and along sented included Rhododendron nudiflorum, Vacci- with it are varyingamountsof red oak, whiteoak, nium stamineum and Kalmia angustifolium. As in the black oak, red maple, and hickory(Fig. 13). The previousstands,erieads contributeover 90% to the shrub layer is typicallyericaceouswith Gaylussacia shrubcover. baccata, Vaccinium angustifoliumr, and Vaccinium The herbaceouslayer is sporadicallydeveloped. A vacillans mostcommon(Fig. 14). Those herbswhich total of 29 herbs occupy less than 7% of the area. contributethe greatest cover are Aralia nudicaulis PINE
SHRUB1
SORUS
PINE
C ESTNUI WNITZ MIXED
OAK
OAK
OAK
OAK
NlCKORY WA*DWO1
OAK
NO. NARDWOODS SWAMP
waWTN
oADwOOo
"""MLDO
QUERCUS ILICIFOLEA_
_
VAcciNIUM ANGUSlTIOLIUM
PYRUS KhELANOCARPA
AM-LANCHIER STOLIONWEIRA
DIERVILLA LIONICE!A NEMOPANTHUS MUCRONATA COMPTONIA PFREGRINA
GAYLUSSACIA BACCATA __ VA CINIUM VACILLANS*
KALMIA LATIFOLIA. 1 K, ANGUSTIFOLIA
S
CORNUS RUGOSA_
RHODODENDRON NIUDIFLIORUM VITIS AESTIVALIS
C.ANOTHUS AM..ICANUS
VACCINIUM STAMINEUM RURUS SPP LYONIA LIGUSTRINA
CORNUS ALTERNIFOLIA_ CORYLUS CORNUTA_
_.
ILEX MONTANA
SMILAX ROTUNDIFOLLA LINDERA BENZOIN-
_
VIBURNU-MACERIFOLIUM PARTHENOCISSUS QUINQUEFOLIA;
_
_ _1_1
H MAMELIS VIRGINLANA
RHODODENDRON MtAXIMUM VIBURNUMLENTAGO_ VRECOGNITUM
ILEX VERTICILLATA
VACCINIUMCORYMBOSUM
C RNUS R CEMOSA_ RHUSRADICANSl;
Aralia
L
__
_
_
nudicaulis, Carex pensylvanica, Pteridium
aquilinumare the only species representingover 1% cover. Those presentin all stands are Aralia nudi-
caulis, Uvularia sessilifolia, Lysimachia quadrifolia, and Hypoxis hirsuta.
The mosses are restrictedto rocky outcrops and the bases of tree trunks. Betweenthe rock outcrops
Polytrichum ohioense, Leucobryum glaucum and Dicranuim scoparium are most important. Thelia asprella, Leucobryum glaucutmand occasionally Rhodobryuimroseumcover the bases of tree trunks,espe-
cially those of white oak. wherethe On the lower slopes of the conglomerate soils are more continuous,red oak and hickoryare associates with the chestnutoak (10). Together these species with the chestnutoak contribute80% of the totaltree cover. Otherassociatesof minorimportanceare red maple,black oak, sweetbirch,black cherry,butternut (Juglans cinerea) and red ash (Fraxinus pensylvanica).
On these better sites the
and Pteridium aquilinum (Fig. 15). WHITE
OAK-HICKORY
COMMUNITY
is characteristic The whiteoak-hickorycommunity in the wide valley floorsalong Big Flat Brook, on the moisterupland sites of the High Falls Formation (Fig. 16), and occasionallyin the later stages of old fieldsuccession. Two stands representingthis communitywere studied in detail, one (No. 11) located near Big Flat Brook in the southernportionof the park, the other (No. 12) on a moistsoutheastfacing slope above Big Flat Brook (Fig. 1). In bothstandsthe soil is rocky. A thinmorhumus overliesa grayish-blackA horizonwhichblends into a yellow-brown layer, the B horizon. The soil pH is approximately5. the forestconsistsof two more In this community, or less distincttree layers,a shrublayer,and sparse herb and moss layers (Fig. 13, 14, 15). In the tree layer,white oak contributesabout 32% and hickory
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April, 1958
PAST AND PRESENT VEGETATION OF HIGH POINT STATE PARK
139 .
*
*
.,~~~~~~Ir
-I.
-~~~~~~~~~~~~
0
FIG. 15. Herb coverand frequency for forestcornmunlities.Lengthof bar represents % covercontributed by each speciesand the widthof the bar represents the frequencyof occurrence based on the total numberof herbquadratsin all standssampledof each community. along road to community FIG. 16. Whiteoak-hickory The (Xr) indicatesthat the specieswas presentin the part of the park. community but not occurringin the quadrats. Space Life Campin southern representsthe amountof unoccupiedarea in the herbaceouslayer. the cover, Harnamelisvirginiana17% of the cover and Viburnumacerifolium16%. A few otherspresabout 15% of the total tree cover (Fig. 16). The ent are listed in figure14. importanceof hickoryvaries considerablythroughout In the herbaceouslayer a verysmall amount,about the community.Otherassociates are black oak, red Ao of the area, is occupied. Of the 54 species presmaple and beech (Fagu8 granclifolia). Sugar maple ent, none contributesmore than 1% cover. Those (Acer 8acchbarum) and yellow birch frequently most frequentlyfound include Uvularia perfoliata, border the streams where they flow throughthis U. sessilifolia,Potentilla simplex, Solidago caesia, community. Sugar maple and white ash are most Aralia nudicaulisand Maianthemum. canadeuse (Fig. abundant in the smaller 1-3.9 in. size class, whereas 15). whiteoak, hickoryand black oak are most common The moss layer is poorly developed. On and bein the larger size classes. The trees are tall and twveen the rocks,Dicranumscoparium,Thuidiumdelistraight. Thereis no evidenceof sproutorigin. The catulumand Polytrichumohioensewere estimatedaS heightand diameterof a few of the larger treesare contributing a coverof 4%. whiteoak, 26.7 in. d. b. h. (85 ft.), hickory(Carga MIXED OAK-HARDWOODSCOMMUNITY glabra) 20.9 in. (80 ft.) and black oak 15.8 in. (74 ft.) (Stand 12). Chestnut,hickory,and white oak The broad sloping valleys betweenthe ridges and logs are found on the forestfloorin various stages hills are occupied by the mixed oak-hardwoodscomof decomposition(Stand 12). Red maple,juneberry, munity(Fig. 1). A standstudiedin detail representdogwood,whiteash, and youngtreesof the dominant ing this communityis located above Saw Mill Pond species comprisethe understory.Nearer the streams on a gentleeast facingslope (Fig. 1, Stand 13). The hornbeamand hop hornbeamincreasein importance soil thereis relativelyrockywithan organiclayer of (Stand 11). The dominanttrees are reproducing 1-3 em. This organiclayer appears to be rapidlyinthemselves. Both seedlings and saplings of sugar corporatedinto the underlyingsoil. Moist spots are maple, birch,and whiteash are common(Stand 11). common,whose degree of mesophytism is dependent About ',3 of the shrublayer is occupied by shrubs. to some degree upon the drainage from the upper Corylue cornutat contributesan average of 24% of slopes.
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140 140
WILLIAM
A.
Ecological Monographs 23, No. 2 ~~~~~~~~~~~~~~~~~~~~~~
NIERING
cimin thicknesscoveringthe rockysurface. Where soil is presentamong the rocks,it consistsof a fine clay loam witha pH of about 5.0. lightyellow-brown The forestis composedof two distincttree layers and sparse shruband herb layers. The tree canopy is nearlycompletewithveryfew openings. Most of the trees are tall and straight. In the upper story are sugar maple and its associates-basswood, red oak, beech,yellow birch,white oak and sweet birch (Fig. 13). Locally beech and sugar maple are most important.Large sporadic whitepine toweringabove the othertreesare sometimespresent. Chestnutwas formerlyan importantelementas indicatedby the huge logs on the forest floor. Yellow birch is restrictedprimarilyto the streammargins. The dominant trees vary from75 to 94 ft. in height; for example one sugar maple 29 in. d.b.h. measured94 ft., a basswood12.4 in. 73 ft.; and a whiteash 19.4 in., 75 ft. Hornbeam and hop hornbeamwith young sugar maples constitutea sub-canopy layer. The trees differin their reproductivepotential. Sugar maple is abundantlyreproducingitself (Fig. 17). Small plants 1-2 ft. in heightare most common,a total of 784 such individualswere recordedin 400 sq. meters. Yellow birchseedlingsare abundant,but saplings are absent. Basswood reproductionis relatively sparse, and beech is reproducingchieflyby root sprouts. White ash reproductionis important in both seedlingand sapling stages. The shrub layer is sparse, occupyingonly 14% of the total area (Fig. 14). To this14% cover,Lindera beuzoin contributes46%. The other 5 species in order of decreasingimportanceare Hamamelis virNORTHERN HARDWOODS COMMUNITIES giniana with24% of the cover,Viburnumacerifolium NORTHERN HARDWOODS WITHOUT HEMLOCK with 18%, Parthenocissus quinquefolia with 7%, The northernhardwoodscommunitywithouthem- Rhus radicans with 3%, and Corylus cornuta with lock is very restrictedin its distributionin the park 92. The herbaceouslayer is also sparsely developed, (Fig. 1.) The beststand is locatedin a narrowrockv valley in the southernportionof the park (Stand 16, only 9% of the area being occupied. The two most Fig. 17). The ground consists mostlyof rounded importantspecies are Arisaema sp. with 3% cover boulders which are often covered with an organic and Symplocarpusfoetiduswith1%. Of the thirtylayer. In places on the valley floor,the bouldersare fiveothersthat occur, those most frequentlypresent exposed, since the organic material is frequently are listed in figure15. Those present but not rewashed away by spring flood waters. Where not cordedin the quadrats are Allium tricoccum,Adiansubjected to such flooding,the humus layer is 2-3 turnpedatum,Osmunda cinnamomea,0. claytoniana and Dryopterisnoveboracensis. The mosslaver covers2% of the area. Those most commonon the rocks are Thuidiumdelicatulum,Dicranumfulvumand Grimmiaapocarpa. The forest consists of two distinct tree layers, a shrub layer and sparse herb and moss layers. It to designatea single dominantsince is oftendifficult many tree species contributesignificantlyto the canopy. Those representedcontributethe following cover: whiteoak 21%, red oak 22%, white ash 17%, sugar maple 12% and red maple 9%o. Other trees interspersedamong these are shagbarkhickory (Carya ovata), chestnutoak, scarlet oak, black gum and beech. Juneberryand young trees of the main canopy comprisethe understory. The tree layer is continuous,wvithno gaps in the canopy (Fig. 13). The larger trees are 14-16 in. d. b. h. and attain a heightof 65 ft. The dominanttreesare reproducing themselves. The white and red oak, while most importantin the tree layer, are less successfulin the reproductivestages. Red maple, sugar maple and whiteash are reproducingprolifically. The sparsely developed shrub layer contributes a totalof only24% cover. Amongthoserepresented, Corylus cornutacontributes42% of the cover,Hamamelisvirginiana35%, and Viburnumacerifolium 16%. Othersof less importanceare listed in figure 14. In the herb layer,38 species contribute13% cover mostto the cover are (Fig. 15). Those contributing Adiantum pedatum with 3% and Mitchella repens with 1%. Others commonlypresent are listed in figure15. Mosses cover only 1% of the area. They occur on the moist exposed soil and between the outcroppingrocks. Thuidiumdelicatulumand Polytrichumohioenseare most common.
NORTHERN
FIG.
17.
Northern
hardwoods
without
hemlock.
Forest floor densely coveled w^itllsugar maple saplings. ( Stand 16.)
HARDWOODS
WITH
HEMLOCK
The northernhardwoodscommunitywith hemlock present is also restrictedin its distribution.It occupies steep boulderyravines (Stand 17) and low islands betweenstreams(Stand 18). One stand studpart of ied in detail is located in the northwestern the park and anotheris situated3/4mi. southof Saw Mill Pond. The soil of the two sites varies considerably. On the steep,rocky,ravines,it consistschieflyof organic material which accumulates between the boulders. However, some of the accumulatinghumus is con-
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PAST AND PRESENT
April, 1953
OF HIGH
VEGETATION
POINT
STATE
141
PARK
stantlyremovedby floodwatersin the narrowravines forest,as well as in abandoned fieldsand former and by erosionon the steep slopes. pastureland. An exceptionto this occurs just outsidethe park, The shrub layer of this communityis relatively east of Lake Marcia. There a stand lies at the base open, shrubs occupyingnot more than 25% of the of the southeastescarpmenton a northfacing slope, area, usually less. Those shrubs which most frewherethe conglomerategives way to the underlying quentlyoccur are Hamamelis virginiana and Viburshale. At this point a streamhas cut a deep ravine. ntunacerifolium (Fig. 14). Rhododendroit maximum The soil in this hemlockstand is derivedfrom the is only locally important. shale and is relativelyfreeof rocks. Wherethe comHerbs are sporadic occupyingabout 6% of the munityoccurs on islands betweenstreams,the soil is area. Those which are found most frequentlyina rockyclay loam witha podzol profile. An organic clude Aster divaricatus, Maianthemum canadense, layer overliesan ash gray A horizon(pH 4.2) below Trientalis borealis and Mitchlella repens (Fig. 15). whichis a yellow-brown B horizon (pH 4.8). Lycopodium lucidulum was present but it was not In the rocky ravines, hemlockand yellow birch foundin the quadrats. are the importantspecies, contributing57% of the Mosses cover 9% of the area in the moss layer, total cover of the forestcanopy (Stand 17). Other the rest being either litter or bare rock. Those main canopy associates in order of dominanceare present are Hylocomium splendens, Dicranum scosweetbirch,whiteash, red maple, sugar maple,bass- pariumn, Climacium americanum, Mnium affine, M. wood,tulip tree,and whitepine (Fig. 13). Hemlock punctatum, and Trichocolea tomentella. In the is more importanton the island sites and on the streamflowingthroughone hemlockstand (17), are shale slope (Fig. 20). Here, along with the above matsof Eurhynchium rusciforme and Sematophyllum associates, white oak. chestnutoak, and black gum carolinianum coveringthe rock surfaces. may be present. The larger trees vary from12 to POLLEN ANALYSIS 32.5 in. d. b. h. The larger hemlocksare 80-100 In general, the pollen record reveals an initial years old. The average height of the forest is 70 forest succeeded by a period of ft. An understorytree layer is poorly developed, spruce-pine-fir-oak and thena briefperiod of pine domiwith striped maple, hop hornbeamand transgres- spruce-pine-fir sives of the dominantspecies contributing the most. nance. This was followedby a deciduouscomplexin All trees are reproducing,althoughoak is least suc- which oak predominated. During the latter, two cessful. Hemlockand birchseedlingsare mostabun- periodsof hemlockmaximawere separatedby an indant. Occasionally hemlockseedlings are so abun- terveninghickorymaximum. Although the two bogs show the same general dant on the forestfloorthat they produce a green tinge. The seedlings of birch and hemlockoccur trends, they do not correlate in the lower levels chieflyon hemlocklitterand on rottenlogs. There (Fig. 18, 19). Some of the recordsrevealedin the is some evidence that this communitymay in time bottomlevels of the Pine Bog profileare absent in occupy a greaterarea, since in at least threestands, the profileobtainedfromCedar Bog. What is espehemlocksaplings are present not only within the cially conspicuousin the Cedar Bog profile,is the stand, but also in the surroundingchestnutoak shortnessof the spruce and firrecordcomparedwith
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P ERCEN TAG E 18. Cedar Bog pollen spectrum.Length of the bar represents% of total tree pollen. For example, Pinus represents18%o at the surface(s) and 68% at the 19 ft. level. Chamaecyparis (cedar) is founldonly at the surface where it represents10% of the total tree pollen.
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142
WIL~ WILLIAM
.NEIGEcological A. NIERING
Monographs 23, No.2 ~~~~~Vol.
4 W
LU
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W
a: O
S .La5y
D
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0.
<-p
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) >
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26 27 29 30 31 32 33
34 35 36 37 368_
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FIG. 19. Pine Bog pollen spectrum.Length of the bar represents%~, of total tree pollen. For example, Pinus represents7% at the surface(s) and 46%/at 40 ft. level. No pollen was found in the 39 ft. sample.
that of Pine Bog. In Cedar Bog, the 20 ft. level is and the reappearance of fir. Beech increased also comparableto the 33 ft. level in Pine Bog. The rec- during the hemlockmaxima. Surface samples of ord found in the bottom7 ft. fromPine Bog is ab- Pine Bog, which presumablyreflectthe presentday sent fromthe Cedar Bog profile. Failure to findthe conditions,show oak and birchto be mostimportant deepest part of Cedar Bog may be the explanation with a slightdeclinein pine and a rise in hemlock. for the incompleteprofilethere (Potzger & Wilson In contrast,oak is much less importantin Cedar 1944). Althoughsamples were obtainedto a depth Bog, where it is accompanied by an increase in of 30 ft. in Cedar Bog, those from 21-30 ft. were hemlock,birch,maple and pine along with appearsterile, except for a few pine and spruce pollen ance of Nyssa and Chamaecyparis. grains at about the 25 ft. level. Small quantities of non-treepollen were found In the bottom levels (37-40 ft.) of Pine Bog, throughoutthe profile. In the 1-4 ft. levels of Pine spruce, fir,pine and deciduous species occur. This Bog, Compositae,especiallyAmbrosia,representsup is followedby a spruce maximumin whichthe de- to 20% and in the lower levels sedges and grasses ciduous elementis practicallyabsent. A pine period, were present. In the top levels of Cedar Bog, whichhas been often shown to be very pronounced Chenopodiumand Rumex occur. in eastern North America (Potzger & Otto 1943; DISCUSSION Deevey 1939, 1943), if representedin these profiles was of short duration being recorded only in the That climate controls vegetation has long been 32 ft. level in Pine Bog and the 19 ft. level in Cedar recognized,the major vegetationalpatternsbeing deBog. This is succeededby a long oak-hemlock-pine the regional climates (Tansley 1949; and mixed broadleaved period during which two terminedby & Clements 1938). Withinany region,one Weaver maxima hemlockmaximaoccurred. The firstof these (around the 15 ft. and 11 ft. level) is accompanied encountersplant communitieswhose presence and are determinedby local climateand edaby a decreasein pine and oak. Betweenthem,there distribution occurred a period in which there was a decrease phic conditions,as for example a ravine compared in hemlockand an unmistakableincreasein hickory. with an exposed ridge, or a site with a thin soil The second hemlockmaximum(5 ft. and 2 ft.) is mantleover rock comparedwith a deep fertilesoil. accompaniedby an increasein chestnutand spruce, It is evident,therefore,that superimposedupon the
This content downloaded from 131.125.1.217 on Thu, 18 Apr 2013 11:13:59 AM All use subject to JSTOR Terms and Conditions
April, 1953
PAST
AND PRESENT
VEGETATION
OF HIGH
POINT
STATE
PARK
143
cially oak, rapidly became importantand have pers'sted until the present. This change is suggestive of a warmerclimate. Associated with these deciduous trees there appeared the hemlock. Its history is accented by two periods of greater importance. These two periodsmay be interpretedas timesof increased nosture (Potter 1947). This interpretation by a correlateddecrease in is furtherstrengthened hickoryat the hemlockmaxima. Betweenthe hemlock maxima a rise in hickoryis indicativeof drier and PAST VEGETATION possibly warmer conditions. This level probably One mightexpect, rs Transeau (1903) did. that markswhat has been knownas the last xerothermic tundra vegetationfirstoccupied the area after the period which had widespread effectsupon eastern ice melted. However, no evidence for such was North America (Sears 1942; Raup 1937; Transeau found, althoughDeevey (1951) reports pollen evi- 1935; Gleason 1923). From samples obtained in denceof tundravegetationbeingfoundin Maine and Connecticutthe date of the hickorymaximumhas possiblyin Connecticutbogs. The beginningof the been tentativelyset by the radiocarbonmethodat recordin Pine Bog shows the presenceof deciduous about 1800 ? 500 years ago (Flint & Deevey 1951). speciesaccompaniedby pine and a low representation The second hemlockmaximumwas accompaniedby of spruce. This suggestsa period of moderatecli- an increasein chestnutand a declinein hickorywhich mate at the start of the record. Since the area lies togetherindicate a returnto more mesic conditions. withinthe Cary moraine and about 100 mi. south Evidencefor such a shiftis strengthened by thereapof the Mlankatomoraine (P. Ma(eClintbck, pers nal pearance of spruce and fir. comminunication) it suggeststhat considerableclimatic And indicationof the historicalperiodis suggested ameliorationoccurredfollowingthe Cary. This was towardthe top of the profiles. In Pine Bog the local followedby a disappearance of the deciduousforest influenceof a developingbog forestmay to some exspecies and a spruce dominancewhich may he cor- tent mask the true picture of upland conditions. It relatedwith the readvanceof the Mankato substage. is possible that the increase in pine near the top Deevey (1939, 1943) found a similar fluctuationin may be evidenceof such an influence. The appellaConnecticutwhich he correlatedwith recessionand tion Pine Swamp along with the presence of white advanceof Early and Late Mankatosubstages. How- pine and dead snags in the bog today would suggest ever,fromrecentstudiesin Maine,Deevey (1951) sug- that pine was probably more importantin recent gests a different interpretation.In these studies he time than at present. The pronouncedrise in birch eonsideredthe rise in spruce as recordingthe actual accompanied by a decline in hemlocksuggests the invasion of spruce into the region, but below the temporarydominanceof gray birch as a result of spruce maximum he considers the tree pollen firesand man's activity. wind-horne froma distance. However,this does not The surfacelevels of Pine Bog reflectthe present seemto he a likelyinterpretation at High Point State upland vegetationdominatedby oak and otherdeciduPark since thereis a paucityof non-treepollen. ous species. Hemlock has increasedslightly,but its Pine becomes importantearly in the profile,as still restricteddistributionin the most mesic sites is comparedwith the vegetationalsequences illustrated reflectedin the surfacelayers. In contrastin Cedar by profilesin the midwest;a phenomenonthat may Bog, theupland forestis partlymaskedby themature he the resultof climateunder more maritimecondi- hog forestcoveringmuch of the bog today. This is tions (Potzger & Otto 1943). Climatically the shownby the increase in hemlock,birch,and maple periodwas cool and mnost.The subse- along withthe appearance of gum and cedar. spruiee-pine-fir quent brief period of pine dominancesuggestsdrier An interestingcorrelationresults when Pine Bog conditions(Sears 1948). It is of much shorterdu- is interpretedin terms of Sears (1948) suggested ration than the pine period recordedby most bogs correlationbetweennortheasternpollen profilesand studiedin the Northeast:in New Jersey (Potzger & climaticfluctuations since early Wisconsintime. The Otto1943), in Connecticut(Deevey 1939, 1943,1951) substageadvancesmay be correlatedwithan increase and in New Hampshire (Krauss & Kent 1944). How- in hemlockor spruce,and retreatswitha rise in pine, ever, it is comparablewith Deevey's Linsley Pond or oak and hickory,concurrentwith a decrease in (L-9 and L-10) profiles. Radiocarbonstudies show hemlockor spruce. Beginning at the top of the the time of attainmentof the pine period in North profile,the low percentageof hemlockand higher America to vary roughly with latitude. In West percentageof oak may reflectthe retreatof the AlVirgin-a it was attained about 9,000 years ago, in pine-Alaskan, although the historical period with Connecticut8,000 and in Maine 6,000 years (Flint man's influencemay be equally or more important. & Deevey 1951). Since these bogs are located rela- The increased importanceof hemlockbetween the tively near Connecticutthe pine period probably 3-9 ft. levels may be correlatedwith the Alpineoccurredaround 8,000 years ago in northernNew Alaskan advance. The retreat of the Cochrane is Jersey. suggestedat the hemlockminimumand hickorymaxiFollowingthe pine period deciduousspecies. espe- mum (10-11); and its advance may be correlated pervadinginfluenceof the regional climateare both physiographicand edaphic conditionswhich locally influencethe vegetationalpattern. Hence, in any regionalvegetationalpattern,therewill be a matrix of vegetationin whichthe regionalclimateappears to have the principal control,but within which pronouncedlocal variationsoccur. The wholepatternis subject to change as changes in the regionalclimate occur.
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144
WILLIAm NIERING WILLIAM A A.NIERING
withthe precedingrise in hemlock,reachinga maximum at the 15 ft. level. The previous decline in hemlockis suggestiveof theAgassiz-Campbellretreat, and it is precededby a slightincreasewhichmay be correlativewith the advance of Agassiz-Campbell. The brief pine period may be correlatedwith the retreatof the Mankato. The Mankato advance is suggestedby the spruce maximum. The bottom sediments represent the period after the retreat of the Cary, which may have been accompaniedby pronouncedclimaticameliorationindicatedby the presenceof oak. If these correlationsare valid thenthispollen profilesupports MacClintock'sbeliefthat the bog has been developed on drift of Cary age (MacClintock,personal communication). PRESENT
VEGETATION-HISTORY
Ecological Monographs 23, No.2 ~~~~~~Vol.
cal of cut-overoak woods, in the evidenceof fires, and in the secondarysuccession patternsfrom old fieldsthroughgray birchto as yet unstabilizedforest. VEGETATIONALPATTERN
The dominantcommunity is the chestnutoak with its characteristicericaceous shrub layer. It forms a matrix in which all the other communitiesare found (Fig. 1, 20). It is relativelystable on most sites,but on some of the lowerslopes it is yieldingto the northernhardwoods. Since the disappearanceof chestnut,natural replacementby oaks has followed. In Stokes State Forest, Korstian & Stickel (1927) noticedthis trend. The typical communityof the rocky ridge top is the pitch pine-scruboak community(Fig. 20, 21). Fires in the past have probablyaided in perpetuating this communityby stimulatingsprout growth of scrub oak and furnishingfavorable conditionsfor pine reproduction(Lutz 1934; Conard 1935; Little & Moore 1949). In fact, the vegetationresembles that foundin parts of the Pine Barrens of southern New Jersey where frequentfires occur. With the exclusion of fire in the pitch pine-scruboak communityon the ridge top, the evidenceindicates that both pines and arborescentoaks have great difficulty becomingestablished,and that shrubsmay continue to dominateover considerableportions. Those pines
With the arrival of man, new influenceswere superimposedupon thosepreviouslycontrolling the vegetationalpattern. Man's activitiesthat affectedvegetation were cuttingthe forests,carelessnesswith fire and cultivatingthe land. These activities resulted generallyin more xeric conditionsand thereforea retrogression in the vegetational development (Nichols1913a). The initial anthropeicinfluenceupon the region began with the Indian, who depended mainly upon fruitsand animals for his existence. He is known to have frequentlyset fires either accidentallyor PITCH PINE *-------CHESTNUT OAKpurposely to facilitate hunting (Little 1946, VerRED OAK SCARLET OAK---meule, et al 1900). Later as whiteman enteredthe HICKORIES region he gradually cleared the areas suitable for HEMLOCK SPACE 1700 agriculture. Later, industrygrew up and this was accompaniedby increasinglyintensiveharvestingof o00 wood from the remainingforestedareas. The dependenceof industryupon forestresourcesreached its maximumduringthe middle1800's. Large quanSHAWANGUJNK 400 titiesof wood were requiredto supply iron furnaces scatteredthroughoutnorthernNew Jersey. During CONGLOMERATE 1300 this period trees 20-25 years of age were cut and utilized as charcoal. Much wood was also used for manyotherpurposessuchas railroadties,hoop poles, 0 200 l00 500 o00 400 and domesticfuels. Around 1850 when anthracite M. SE blast furnaces were introduced,the utilization of FIG. 20. Transect repiesenting thee cover (Fig. 1, forestresourcesbegan to decrease (Vermeule,et al Tranisect B). The distance of each line representing 1900). However, by this time portionsof the ad- the species above the topographic pi ofile indicates the % of tree cover representedby the species at different jacent New Jersey Highlands were completelyde- points along the transect line. For example, on the nuded of forest vegetation. Along with the inten- ridge top (0-50) pitch pine contributes88% of the total sive cutting were severe forest fires. During the tree cover and open space in the tree canopy is at its special surveyof forestsfrom1894-1899,Vermeule, greatest (70% ). Other species are present but altoet al (1900) pointed out that pitch pine as well as gethercontributeonly 12% of the cover and arc omitted here. On the slopes, chestnutoak contributes62% cover, mixedconiferousand deciduousforestsof the Kitta- reaching a maximum of 71% (190), followed by aln tinny Mountains sufferedconsiderably. Although abrupt decrease to 25%/ on the lower slope (360-460). a permanentorganizationto controlfireswas estab- Chestnut oak on the lower slope is accompanied by aplished as a result of this survey,frequentfireshave proximatelyequal percentages of hickoryand red oak. oak is restricted primarily to the upper slopes since plagued the region. Present residentsrecall Scarlet (60-2.50). Numbers at the bottom refer to horizontal severallarge firesin morerecenttime,one or moreof distance in metersand at right,refer to altitude in feet whichburnedover large areas along the major ridge above mean sea level. Glacial drift overlyingthe geological formationsis shown by stippled area with larger withinthe park. dots. Talus slopes are represented by small angular Today the effectsof man's activitiesare evident blocks. The Martinsburg shale is showvnat right by everywhere:in the prevalenceof sproutgrowthtypi- olblifle liues.
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April, 1953
PAST
AND PRESENT
VEGETATION
OF HIGH
POINT
STATE
PARK
145
Where land has been cultivatedat some time in the recent past, gray birch is now common. Raup (1937) found the same situationin the Black Rock Forest of New York. Red cedar also occurs,but is not abundant. It is primarilyrestrictedto the shale. In southernNew England, Lutz (1928) and Raup (1940) describe both red cedar and gray birch in secondarysucceession. Bard (1952) found red cedar abundantand littlegray birch on the Triassic shales of thePiedmontof New Jersey. The birchand cedar, which are relativelyshort lived, are replaced by an oak forest. Since the recessionof the ice sheet,primarysuccessionhas been continuously occurringon thescoured ledges of the ridge and in the glacial lakes. In fact, thereis still bare rock to be coveredand waterto be filledin withvegetation. The earliersuccessionalpatternswere probablynot unlikethose operativetoday. The species concernedmay well have been different; but the earlierthe stage in the succession,presumably the morenearlythe presentvegetationresemblesthat FIG. 21. Transeet representing tree cover (F a. 1, Transect A). See Figure 20 for guide in interpretation. of early post-glacialtime. Plants that suggest this Cedar Bog deposits are represented by parallel lines. are lichens,mosses,and certainfloweringplants such PITCH PINE CHESTNUT OAK RED OAK HICKORIES RED MAPLE WHITE OAK SWEET BIRCH WHITE BIRCH HEMLOCK YELLOW BIRCH SO-WHITE CEDAR SLACK SPRUCE SPACE
..
NW
...........
SE
Break in line across Bog (210-360) indicates that only a portion of bog was sampled. The letter (R) on either side of the bog indicates location of road.
as Deschampsia flexuosa and Potentilla tridentata.
arborescent oaks and the oaks are reproducing prolifically.compared to the pine. Evidence of greater importance of pine in the past is indicated by rotten pine logs on the forest floor, by the occurrence of a podzol soil profile and by sporadic clumps of scrub oak. Very similar to this are late successional stages in the Pine Barrens of New Jersey (Buell & Cantlon 1950). In the absence of further pyric and anthropeic influences, the pine will form only a small part of the stands if present at all (Little & Moore 1949). The white oak-hickory community occurs on the valley floors or in more mesic upland sites (Fig. 21, 22, 23). The presence of chestnut logs in some stands indicates that where hickory is present today both species, along with white oak once shared dominance. At present this community shows a successional trend toward the northern hardwoods. There is an increasing importance of the northern hardwoods in the smaller size classes. The mixed oak-hardwoods community in the same way is yielding to the northern hardwoods. The established northern hardwoods are at their maximum development in deep narrow valleys (Fig. 20, 23). Further aggressiveness by the more mesie species is evidenced by the young hemlock which is in the adjacent chestnut oak community.
meda glaucophylla, black spruce, tamarack and asso-
The processof primarysuccessionis muchthe same for all rock surfacesalthoughthe ultimatevegetation which becomes established depends upon the expresent are primarily in the open early stages of posure. On the ridge tops it leads to pitch pineprimary succession. This community will probably scrub oak or possibly to chestnutoak. In contrast, continue to be represented as long as primary vegeon the northwestslopes are northernspecies such tational succession is still in the process of covering as white pine, white birch, sweet birch, mountain the bare rocky ledges, and as long as fires are allowed ash, striped maple, and mountainmaple associated to sweep across the ridge. with chestnutoak. In contrast, the pine-oak community as it occurs The early stages of bog successionat High Point, today, on the deeper soils derived from glacial drift, as in the rock succession,reflectthe early boreal enis an advanced successional stage approaching oak vironmentshown to have existed in the area by the forest. Sporadic pines frequently tower above the pollen record. For example,species such as Androciated vegetationappear as relictsaround whichmore southernspecies have becomeestablishedas Transeau (1903) long ago pointed out. He believed that the presence of either a coniferousbog or broadleaved swamp vegetationdepends upon the period in postglacial timewhena suitablehabitatbecame available. Althoughboth occupy the same kind of habitat today, the coniferousbog has been gradually losing out. Only where the conifer vegetation has been reasonablyundisturbedhas it persisted. Elsewhere, the swamp hardwoods have replaced it, and even withoutdisturbancethe successionappears to lead to a swamp hardwoodforestwithhemlock. The occurrenceof southernwhite cedar in Cedar Bog is of particular interest. Its invasion into the area may have occurredduring the last xerothermic period or hickorymaximum. Several investigators have found evidenceto indicate that cedar may have been more importantin the recent past (Bartlett 1909; Nichols 1913b; Raup 1937; Heusser 1949). Today it is decreasingand will ultimatelygive way to the more shade toleranthemlockand red maple (Little 1950). The broadleaved swamp forest of more recent origin is relatively stable, with little
This content downloaded from 131.125.1.217 on Thu, 18 Apr 2013 11:13:59 AM All use subject to JSTOR Terms and Conditions
146
A.
WILLIAM
Ecological Monographs Vol. 2 3, No. 2
NIERINO
CHESTNUT OAK RED OAK HICKORIES RED MAPLE WHITE OAK SWEET BIRCH WHITEPINE WHITEASH SUGAR MAPLE SPACE
~~~~~~~~~~~~~0 ~ ~ ~
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CHESTNUT OAK WHITE OAK RED MAPLE SUGAR MAPLE BEECH SPACE
.
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FIG. 23. Transects representingtree cover (Fig. 1 Upper transectE, Lower D). See Figure 20 for sample explanation.
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evidenceof changeexcept whereaccumulationof soil has resultedin more mesicconditions. It is interesting to note that relict species whose range is more continuouseither furthernorthor south are found in both xerarch and hydrarchprimarysuccessions, which offerless competitionthan other habitats. Today the mosaic of vegetationin High Point State Park is composed of plant communitiessome of whichare relativelystable,while othersshow successional tendencies. In recenttimethe activitiesof man have changed the forest from one which was more mesic, having a larger representationof northern hardwoods,to the more xeric types of today. The northernhardwoodsare very restrictedin their distribution;howeverthey are expandinginto the surroundingoak communities, a trend noted by Raup in the Black Rock Forest (Raup 1938). With protectionratherthan destructionmoremesic conditions will develop, although the physiographyand soils will always exert pronouncedinfluenceslimitingthe possible extentof mesophyticdevelopment.
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SUMMARY 1. The post-glacial historyof High Point State Park reveals an initial spruce-pine-fir-oak forest succeededby a period of spruce-pine-fir and then a briefperiod of pine dominance.Finally therewas a prolongeddeciduouscomplexin whichoak predominated. During the latter, two periods of hemlock
This content downloaded from 131.125.1.217 on Thu, 18 Apr 2013 11:13:59 AM All use subject to JSTOR Terms and Conditions
April, 1953
PAST
AND PRESENT
VEGETATION
OF HIGH
POINT
STATE
PARK
147
maximum with an interveninghickory maximum Braun, E. Lucy. 1950. Deciduous forests of eastern frommesic,to xerother- North America. Philadelphia: Blakiston Co. 596 pp. suggestclimaticfluctuations Bray, W. H. 1930. The developmentof the vegetation mic, witha returnto mesic conditions. of New York State. -N. Y. State Col. Forestry. with an asso2. Today a chestnutoak community Tech. Publ. 29. ciated ericaceous shrub layer is the continuouselementof vegetationwithinthe park. In otherwords, Britton, N. L. 1884. On the existence of a peculiar flora on the Kittatinny Mountains of northwestern it forms a matrix in which occur all other comNew Jersey. Torrey Bot. Club Bull. II: 126-128.
munities.
3. The dispersed communitieswithin this matrix Buell, M. F. & J. E. Cantlon. 1950. A study of two communitiesof the New Jersey Pine Barrens and a and theirtypical sites are: the pitch pine-scruboak comparison of methods. Ecology 31: 567-586. on the rocky conglomerateridge tops; the pine-oak Conard, H. S. 1935. The plant associations of central on hills of the High Falls; the whiteoak-hickoryon Long Island. Amer. Midland Nat. 16: 433-516. many of the valley floors,and some of the moist Deevey, E. S., Jr. 1939. Studies oln ConnecticutLake uplands; the mixed oak-hardwoodson the slopes of sediments. Am. Jour. Sei. 237: 691-724. broad valleys betweenthe hills; the northernhard. 1943. Additional pollen analyses from southwoods in the narrowervalleys and deeper ravines; ern New England. Amer. Jour. Sei. 241: 717-752. and of course,early successionalstages of both pri1951. Late-glacial and post-glacial *pollen maryand secondarysuccession. diagrams from Maine. Amer. Jour. Sei. 249: 177-207. 4. Early stages of primaryxerarchand hydrarch Egler, F. E. 1940. Berkshire plateau vegetation,Mass. succession,initiated upon the disappearance of the Ecol. Monog. 10: 145-192. Pleistoceneice, are continuingnow, probably much Fernald, M. L. 1950. Gray's manual of botany, 8th ed. the by is indicated This the same as at theirstart. American Book Co., New York. 1632 pp. part played by several boreal species. Flint, R. F. & E. S. Deevey. 1951. Radiocarbon dating 5. The presence of certain disjunct southernspeof late Pleistocene events. Amn.Jour. Sci. 249: 257cies such as Chamaecyparis thyoides suggestsrelicts 300. period. of the xerothermic Gleason, H. A. 1923. The vegetational history of the 6. Primaryhydrarchsuccessionleads to a swamp Middle West. Annals. Assoc. Amer. Geogr. 12: 39-85' hardwoods forest with hemlock. Disturbance has and Contr. New York Pot. Gard. 242. hastenedthe replacementof the northernconifersby Gordon, R. B. 1940. The primeval forest types of this forest. southwesternNew York. N. Y. State Museum Bull. 7. In secondarysuccessiongray birchplays a dom321. inant role. Red cedar, so importantin southern Grout, A. J. 1940. List of mosses of North America New England and furthersouthin New Jersey,is of northof Mexico. The Bryologist 43: 117-131. minor importance. Andropogonspecies are unim.Heusser, C. J. 1949. A note oln buried cedar logs at portant except on the eastern fringe of the park Secaucus, N. J. Torrey Bot. Club Bull. 76: 305-306. wherethe shale outcrops. N. 1932. A botanical survey of the Tug Hotchkiss, 8. The presentvegetationalpatternis a result of Hill Plateau. N. Y. State Museum. Bull. 287. man's activitiessuperimposedupon climatic,edaphic, and physiographicconditions. On most sites where Hough, A. F. 1936. A climax forest communityon East Tionesta Creek in northwesternPenna. Ecology they occur the chestnutoak, pitch pine-scruboak, 17: 9-28. commuhardwoods northern and hardwoods swamp Hough, A. F. & R. D. Forbes. 1943. The ecology and nitiesare relativelystable. The pine-oak,whiteoaksilvics of forests in high plateaus of Penna. Ecol. show evihickory,mixedoak-hardwoodscommunities Monog. 13: 300-320. oak dence of change. In the morexeric communities Jennings, H., I. L. Owen, I. V. Stone & W. R. Robbers. is replacingpine and in the more mesic communities 1913. Soil survey of the Sussex Area, New Jersey. and the northernhardwoodsare replacing the oaks U. S. Dept. Ag. Bureau of Soils. theirassociates.
Jennings, 0. E. 1939. A contributiontoward plant geographyof westernPenna. Trillia 10: 46-81. C. F. & P. W. Stickel. 1927. The natural Korstian, speclife-form ArchaTd,H. 0. 1953. A comparison of of blight-killedchestnut in the hardwood replacement tra of three New Jersey pitch pine communities. In forests of the northeast. Jour. Ag. Res. 34: 631-648. press. & G. N. Kent. 1944. Analyses and corBard, Gily E. 1952. Secondary succession on the Pied- Krauss, R. W. relation of four New Hampshire bogs. Ohio Jour. mont of New Jersey. Ecol. Monog. 22: 195-215. Sci. 44: 11-17. Bartlett, H. H. 1909. The submarine Chamaecyparis H. B. 1940. The geology of New Jersey. Kummel, bog at Woods Hole, Mass. Rhodora 11: 221-235. Dept. of Conservationand Devel. Geol. Series Bull. 50. chaparral of analysis The statistical 1943. Bauer, H. L. and other plant communities by means of transect Lindeman, R. L. 1941. The developmentalhistory of Cedar Creek Bog, Minn. Am. Mid]. Nat. 25: 101-112. samples. Ecology 24: 45-60. Little, of S. 1946. The effectsof forestfireson the stand Blodgett, F. L. 1910. Ecological plant geography history of New Jersey's pine region. Northeastern Maryland, Midland Zone; upper midland district. Forest. Exp. Sta. Forest ManagementPal-er 2. Md. Weather Serv. Spec. Publ. 3: 221-274. LITERATURE
CITED
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148 148
WILLIAM WILLIAM
A. A. NIERING
Ecological Monographs 23, No. 2 ~~~~~~~~~~~~~~~~~Vol.
1950. Ecology and silviculture of white Raup, H. M. 1937. Recent changes of climate and vegecedar and associated hardwoods of southernNew Jertation in southern New England and adjacent New sey. Yale Univ. School ForestryBull. 56. York. Arnold ArboretumJour. 18: 79-117. Little, S. & E. B. Moore. 1949. The ecological role of . 1938. Botanical studies in Black Rock Forprescribed burns in the pine-oak forests of southern est. Black Rock Forest Bull. 7. New Jersey. Ecology 30: 223-233. 1940. Old field forests of southeasternNew -. Lutz, H. J. 1928. Trends and silviculturalsignificance England. Arnold Arboretum Jour. 21: 266-273. of upland forestsuccessionsin southernNew England. renlund, R. N. 1951. Aquatic vegetation of some Yale Univ. School Forestry Bull. 22. New Jerseyfresh-waterlakes. N. J. Fisheries Survey 1930. The vegetation of Hearts Content,a -. Rpt. l. virgin forest in northwesternPennsylvania. Ecology Salisbury, R. D. 1902. The glacial geology of New II: 1-29. Jersey. N. J. Geol. Survey 5. 802 pp. . 1934. Ecological relations in the pitch pine Sears, P. B. 1930. A record of post-glacial climate in plains of southern New Jersey. Yale Univ. School northernOhio. Ohio Jour. Sci. 30: 205-217. Forestry Bull. 38. - -. 1942. Xerothermic theory. Bot. Rev. 8: Moore, E. B. 1939. Forcst managementin New Jersey. 708-736. Dept. of Conservationand Development,Div. Forests -. 1948. Forest sequence and climatic change and Parks. 54 pp. in northeasternNorth America since early Wisconsin Nichols, G. E. 1913a. The vegetationof ConnecticutII time. Ecology 29: 326-333. Virgin forests. Torreya 13: 199-215. Tansley, A. G. 1949. The British Islands and their . 1913b. Vegetation of ConnecticutI Phytovegetation I. London: Cambridge Univ. Press. 484 geographical-Aspects. Torreya 13: 89-112. Pp. 1935. The hemlock-white pine-northern Transeau, E. N. 1903. On the geographic distribution hardwood region of eastern North America. Ecology and ecological relations of the bog plant societies of 16: 403-421. northernNorth America. Bot. Gaz. 36: 401-420. Costing, H. J. 1942. An ecological analysis of the . 1935. The prairie peninsula. Ecology 16: plant communities of Piedmont, North Carolina. 423-437. Amer. Midland Nat. 28: 1-126. Oosting, H. J. & L. E. Anderson. 1937. The vegetation U. S. Dept. Agr. 1926. Summaries of climatological data by sections. Weather Bur. Bull. W. No. 3. of a barefaced cliff in western North Carolina. Vermeule, C. C., A. Hollick, J. B. Smith, J. Gifford& Ecology 18: 280-292. G. Pinchot. 1900. Annual report of the State GeoloPotter, L. D. 1947. Post-glacial forest sequence of g st for Year 1899. Report on Forests. 327 pp. northcentral Ohio. Ecology 28: 396-417. Potzger, J. E. & J. H. Otto. 1943. Post-glacial forest Waksman, S. A., H. Schulhoff,C. A. Hickman, T. C. Gordon & S. C. Stevens. 1943. The peats of New succession in northernNew Jersey as shown by pollen Jersey and their utilization. N. J. Dept. Conserv. records fromfivebogs. Amer. Jour. Bot. 36: 83-87. & Dcvlpmt. Bull. 55, Part B. Potzger, J. E. & I. T. Wilson. 1944. A study of lake fillingand bog formation. Ind. Acad. Sci. Proc. 54: Weaver, J. E. & F. E. Clements. 1938. Plant ecology. New York: McGraw-Hill Book Co. 601 pp. 79-80.
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