Ecological Monographs, 75(3), 2005, pp. 295±315 ᭧ 2005 by the Ecological Society of America

LANDSCAPE PALEOECOLOGY AND MEGAFAUNAL EXTINCTION IN SOUTHEASTERN NEW YORK STATE

GUY S. ROBINSON,1,4 LIDA PIGOTT BURNEY,2 AND DAVID A. BURNEY3 1Department of Natural Sciences, Fordham College at Lincoln Center, 113 West 60th Street, New York, New York 10023 USA 2The Louis Calder Biological Station, Fordham University, P.O. Box K, Armonk, New York 10504 USA 3Department of Biological Sciences, Fordham University, 441 East Fordham Road, Bronx, New York 10458 USA

Abstract. Stratigraphic palynological analyses of four late Quaternary deposits com- prise a landscape-level study of the patterns and processes of megafaunal extinction in southeastern New York State. Distinctive spores of the dung fungus Sporormiella are used as a proxy for megafaunal biomass, and charcoal particle analysis as a proxy for ®re history. A decline in spore values at all sites is closely followed by a stratigraphic charcoal rise. It is inferred that the regional collapse of a megaherbivory regime was followed by landscape transformation by humans. Correlation with the pollen stratigraphy indicates these devel- opments began many centuries in advance of the Younger Dryas climatic reversal at the end of the Pleistocene. However, throughout the region, the latest bone collagen dates for Mammut are considerably later, suggesting that megaherbivores lasted until the beginning of the Younger Dryas, well after initial population collapse. This evidence is consistent with the interpretation that rapid overkill on the part of humans initiated the extinction process. Landscape transformation and climate change then may have contributed to a cascade of effects that culminated in the demise of all the largest members of North America's mammal fauna. Key words: dung fungi; extinctions; ®re history; human impacts; mastodons; megafauna; mi- croscopic charcoal; paleoecology; palynology; pollen; Sporormiella; Younger Dryas.

INTRODUCTION cally modern humans out of Africa (Martin and Stead- man 1999). Disagreement prevails over what drove the late Pleis- Absolute dates at least broadly constrain the above tocene extinctions in North America (Martin and Stead- North American timeline, which may be resolved by man 1999, Grayson and Meltzer 2003). In spite of clear analyzing microfossil stratigraphies at a landscape lev- associations between an expanding paleolithic (Clovis) el. On a local scale, rates and geographic patterns of culture and the extinct mammal fauna, more conclusive faunal extirpation and cultural activity then can be con- evidence for overkill is lacking. Although the geo- sidered in light of proposed causes of megafaunal ex- chemical record indicates that both developments co- tinction, as outlined in Table 1 (also see Appendix). incide with rapid climate swings, particularly in the Simpson (1953) noted that when a novel invader North Atlantic at ca. 11 000 radiocarbon years before rapidly drives another species to extinction by com- 14 present ( C yr BP) (Alley et al. 2003), the order of petition, chances are against ®nding the two forms at events has been blurred by ¯uctuating levels of at- the same stratigraphic level. Furthermore, rare taxa mospheric 14C. The latter resulted in ``plateaus'' and tend to ``disappear'' from the fossil record before the ``cliffs'' in the radiocarbon calibration curve (Fiedel time of actual extinction (Signor and Lipps 1982). Con- 1999). versely, human remains and artifacts, which similarly However, the North American episode is but one of may be rare initially, would not be expected to ``ap- several pulses of extinction worldwide, ®rst taking pear'' until some time after actual human arrival. We large vertebrates (megafauna) on continental land- have used, therefore, proxy measures that may be more masses late in the Pleistocene and proceeding with is- sensitive to these factors than the conventional fossil land losses for the next 10 millennia. This later phase and archaeological record alone. encompasses historic time, now affecting medium and As well as contributing to a reconstruction of veg- small vertebrates. Each of these episodes of extinction etational and environmental patterns, stratigraphic pol- has occurred in step with initial dispersal of anatomi- len studies here include spore counts of the dung fun- gus, Sporormiella as a measure of large herbivore bio- mass. Sporormiella spore percentages have been found Manuscript received 21 July 2003; revised 28 September 2004; accepted 18 October 2005; ®nal version received 7 January to rise following the introduction of grazing animals 2005. Corresponding Editor: C. C. Labandeira. in historic times. Spores are usually abundant in Pleis- 4 E-mail: [email protected] tocene sediments but scarce or absent for most Holo- 295 296 GUY S. ROBINSON ET AL. Ecological Monographs Vol. 75, No. 3

TABLE 1. Proposed causes of megafaunal extinction in North America.

Hypoth- esis Type Source Rate Pattern Process Comments

H1 climate change Graham and slow? A mosaic vegetation Climate change Lundelius pattern becomes leads to less (1984), Guth- zonal; extinctions hospitable en- rie (1984) follow climate vironments for gradient. certain large species that fail to adapt. H2 environmental King and Saun- slow? Extinction follows Rapid expansion Applies only to insularity ders (1984) as boreal forest of deciduous extinction of recedes north- forest reduces American mas- wards. available habi- todon tat. H3 rapid climate Berger (1991) rapid Follows Younger As H1, but change: the Dryas onset at ea. change is more Younger Dryas 11 000 14CyrBP drastic and de- velops rapidly. H4 blitzkrieg, or Martin (1984) rapid A wave proceeds NaõÈve fauna rap- rapid overkill through a region. idly hunted to extinction. H5 protracted over- Whittington and slow Slow wave, or mo- Initially naõÈve kill Dyke (1984), saic pattern of fauna; overex- Fisher (1997) megafaunal col- ploitation lapse eventually causes col- lapse. H6 predator pit Janzen (1983) rapid Wave through a re- Humans and na- gion tive predators each contrib- ute to col- lapse. H7 second-order Whitney-Smith moder- Pulsed Interactions be- predation (2001) ately tween humans, rapid carnivores, herbivores, and vegeta- tion. H8 three-stage over- Alroy (2001) rapid Pulsed Overkill suf®- kill cient to ex- plain pattern. H9 clovis age Haynes (1991) rapid Severe but tempo- Aridi®cation, drought rary vegetation rapidly ampli- change following fying preda- human arrival, at tion by hu- ea. 11 000 14Cyr mans. BP H10 hypervirulent MacPhee and very rapid Panzootic disease Infectious dis- disease Marx (1997) pattern ease with trans-generic virulence. H11 keystone mega- Owen-Smith not speci- Landscape transfor- Megaherbivores, Fire regime herbivores (1987), Zimov ®ed mation and ®re which main- changes as et al. (1995), follow megafau- tain open for- forests close SchuÈle (1990) nal collapse. est, are re- and fuel loads moved by hu- rise (SchuÈle mans or dis- 1990) ease. H12 great ®re Humbert (1927), rapid Simultaneous Landscape trans- Proposed for Miller et al. throughout large formation by Madagascar; (1999) regions anthropogenic applicable to ®re. Extirpa- North America tion follows loss of forage. H13 synergy Burney (1993a, slow Mosaic pattern of Human and natu- b, 1999), Dia- extinctions ral causes in- mond (1984) teract. August 2005 MEGAFAUNAL EXTINCTION IN NEW YORK 297 cene samples (Davis 1975, 1987, Davis and Shafer 2002). High percentages of Sporormiella on mammoth dung have tied this coprophilous fungus to the extinct megafauna (Davis et al. 1984), and although spores may be found on the dung of rabbits and small rodents, elevated values in modern sediments appear to be ex- clusively associated with a high density of large her- bivores (Davis 1987, Burney et al. 2003). Stratigraphic charcoal studies as a measure of re- gional ®re occurrence may also detect the onset of cul- tural activity in a variety of geographic contexts (Bur- ney 1997). Such work has traced early landscape trans- formation by humans on landmasses bordering the north Atlantic, northern Europe, Australia, South America, Madagascar, and oceanic islands of the Pa- ci®c and Caribbean (Singh et al. 1981, Burney 1987, Burney et al. 1994, 1995, Athens 1997, and references in Grayson 2001). The onset of anthropogenic con¯a- gration often can be distinguished from natural trends in burning: a sharp peak in microscopic charcoal values of at least an order of magnitude is typically followed by one to two millennia of lower concentrations that are nevertheless considerably above background levels. For reasons outlined above, such palynological markers tend to put human activity centuries or sometimes mil- lennia earlier than direct archaeological evidence. This study examines fossil records from the upper Wallkill Valley and the Hudson Lowlands of southern New York State (Fig. 1), an area covered by the Lau- rentide ice sheet at Last Glacial Maximum. Sediment FIG. 1. Location of sites in southeastern New York State, dates from near the terminal moraine, at least 45 km USA. further south, suggest that deglaciation was well un- derway by 18 750 14C yr BP (Muller and Calkin 1993). Terrestrial and meltwater drainage northward became impounded by the retreating glacier, forming a suc- known to be a megafaunal site, has a long sedimentary cession of periglacial lakes in the Wallkill Valley of up record with good temporal resolution. to 90 km2 in area, eventually draining into Glacial Lake Each of the megafaunal sites was examined through Albany (Connally et al. 1989), which would have in- cores and worked as an open excavation (Table 2). A undated the Hudson Lowlands during its earliest stages cone of depression in the water table was maintained (Dineen 1986). It has been estimated that the glacial with a gasoline-powered pump while work was under- margin receded into the upper Hudson Valley by 13 000 way (Burney et al. 2001). Sediment sequences were 14C yr BP, (Connally and Sirkin 1973) but a review of described and samples removed directly from a cleaned available age data suggests the deglacial chronology in vertical face of the excavation trench. An open-sided New York is still not tightly constrained (Muller and bucket auger sampled below the level of open exca- Calkin 1993). Isostatic rebound eventually led to the vation, allowing for descriptions of sediments and sub- southward drainage of Lake Albany into the Atlantic, sampling in the ®eld. Lake sediments were recovered but the Wallkill River continues to ¯ow north and much witha5cmdiameter modi®ed Livingstone piston corer. of its upper reaches are surrounded by extensive muck- In the laboratory, separate subsamples were taken lands. Proximity to the melting glacier has meant that for pollen/charcoal processing, loss-on-ignition (LOI) sedimentary basins of this region had a high rate of determinations using the Dean method (Dean 1974), sediment accumulation encompassing the presumed and 14C dating. Pollen/charcoal subsamples were stored ``extinction window.'' in vials under refrigeration; 14C samples were oven- dried and stored in airtight containers at room tem- METHODS perature. Fossils of extinct Pleistocene megafauna were re- Pollen processing followed standard methods (Faegri covered from well-strati®ed subphreatic sites at Otis- et al. 1989); digests were suspended in glycerol for ville, Hyde Park, and Pawelski Farm (Fig. 1 and Table permanent liquid slide mounts. At least 400 grains of 2). A medium-sized lake, Binnewater Pond, while not tree and shrub pollen were counted per sample using 298 GUY S. ROBINSON ET AL. Ecological Monographs Vol. 75, No. 3

TABLE 2. Characteristics of New York State sites.

Latitude, Elevation Type of Megafaunal Sampling Site General location longitude (m) feature remains method Pawelski Farm Wallkill Valley 41Њ20Ј02Љ N, 122 large wetland Cervalces excavation ``Black Dirt'' 74Њ25Ј45Љ W scotti and bucket auger Binnewater Pond Wallkill Valley 41Њ24Ј36Љ N, 256 lake piston core 74Њ33Ј07Љ W Otisville Wallkill Valley 41Њ28Ј23Љ N, 230 small wetland Mammut excavation 74Њ31Ј10Љ W american- and bucket um auger Hyde Park Mid-Hudson Val- 41Њ46Ј48Љ N, 74 wetland Mammut excavation ley: Hudson 73Њ53Ј38Љ W american- and bucket Lowlands um auger

400ϫ magni®cation; a 1000ϫ (oil immersion) lens was unit contains abundant algal and humic material. A employed for dif®cult determinations. gradual contact at about 11.2 m forms the lower bound- Unless otherwise noted, pollen zones have been as- ary of a nearly uniform black gyttja (Munsell color 5Y signed using stratigraphically constrained cluster anal- 2.5/2) that continues to the sediment-water interface. ysis (CONISS) on the square-root transformed data, The subsampled face of the core revealed close lami- using Edwards and Cavalli-Sforza's chord distance. nations in the lower unit that were most apparent in Only taxa comprising at least 5% of the upland pollen the samples dried for 14C dating. Plant macrofossils are sum in at least one sample have been included in the very sparse, but pollen is abundant, with excellent pres- cluster analysis. Sporormiella spore counts are pre- ervation throughout the sequence. sented as a percentage of the combined aquatic and upland pollen sum. Pollen stratigraphy.Ð Samples for charcoal analysis were treated using a modi®cation of the Winkler nitric acid process (Wink- 1. Zone I.ÐPinus (pine) and Picea (spruce), in that ler 1985) to remove pyrite. Resulting digests were order, dominate the pollen spectrum (Fig. 2). Pollen of washed through 250-␮m sieves to remove macrofossils broadleaf taxa is less important, but Alnus (alder) rises and macroscopic charcoal before hydrogen ¯uoride to 8%, while Salix (willow) and Quercus (oak) each treatment. Calibrated tablets of Lycopodium spores attain 5%. Cyperaceae (sedges) are relatively abundant, were added before the ®nal residues and were sus- up to 15% of the total. Artemisia (sage), Thalictrum pended in glycerine jelly for permanent slide mounts. (meadow rue), and Poaceae (grasses) each attain almost Charcoal residues were counted at 400ϫ magni®- 5%. The date of 22 850 Ϯ 90 14C yr BP (all values are cation using an eyepiece reticule so as to measure the reported as mean Ϯ SD) is on sediment which is not a two-dimensional projected area of individual frag- basal sample and thus does not necessarily mark the ments, reported as ␮m2/cm3. Morphology and texture lower strata of Zone I. of microscopic charcoal fragments are reported here as 2. Zone II.ÐA steady rise and peak in Picea replaces either graminoid (derived from cuticles of grasses and Pinus as the dominant taxon just before the middle of sedges) or amorphous (from woody and broadleaf plant this interval. However, by the upper boundary, Pinus types; Goldberg 1986, Burney 1987). Particles whose steadily recovered, once again outnumbering Picea, smallest dimension exceeded 250 ␮m are recorded as whose values dropped below 10%. Abies (®r) and Betula number of particles/cm3. (birch) rise rapidly to peak at over 10% and 13% re- spectively. Quercus shows a rise and decline, peaking Absolute dates mid-zone at 15%. Herbaceous taxa become less impor- To construct a chronology of latest occurrences, ra- tant toward the upper part of the zone. Five internally diocarbon dates have been obtained on megafaunal re- consistent radiocarbon dates within Zone II range from mains excavated from these sites (Table 2) or alter- 13 210 Ϯ 90 to 11 494 Ϯ 91 (Table 3 and Fig. 2). natively from museum specimens of the same region. 3. Zone III.ÐPinus shows a decrease, while P. stro- Plant macrofossils or sediment samples were dated to bus (white pine) appears fairly abruptly at the bottom help establish the stratigraphic chronology in cores and of the zone, beginning a continuous regional presence excavations. lasting for thousands of years. Picea recovers and re-

RESULTS mains at about 15% before declining sharply to zero at the top. Abies peaks and also drops to zero at the Binnewater pond top while Quercus declines to below 5%. Alnus and Sediment description.ÐA dark greenish-gray humic Betula rise rapidly to 25% and Ͼ30% respectively silty clay (Munsell color 10Y 4/1) forming the lower across Zone III before declining abruptly at the top. August 2005 MEGAFAUNAL EXTINCTION IN NEW YORK 299

FIG. 2. Diagram showing percentage of the total pollen and spores at each depth for major pollen and spore types for Binnewater Pond, New York, USA. Cluster analysis used is constrained incremental sum of squares (CONISS).

4. Zone IVa.ÐA sediment date of 8900 Ϯ 50 14Cyr 5. Zone IVb.Юrst Fagus (beech) and then Carya BP marks the bottom of this unit. Zone IVa thus is (hickory) become important.Tsuga (hemlock) declines characterized by the rapid rise of thermophilous spe- to less than 1%. cies, particularly Quercus which jumps to Ͼ60%, a 6. Zone IVc.ÐTsuga remains at Ͻ1% throughout, level maintained more or less until the present. Betula and appears to be replaced principally by Pinus strobus. and Alnus remain at low levels through the entire zone. Castanea (chestnut), occasionally present in low num- Pinus remains Ͻ10%, dropping Ͻ5% at the top. Ulmus bers since Zones I and II, becomes an important com- (elm) appears, maintaining a steady 3±4% through the ponent of the assemblage toward the top of Zone IVc. late Holocene. A slight rise in Pinus strobus pollen Cyperaceae (sedges) reach values not seen since the occurs at the top of Zone III. late Pleistocene. 300 GUY S. ROBINSON ET AL. Ecological Monographs Vol. 75, No. 3

TABLE 3. Radiometric dates.

Calibrated Stratigraphic Site/ 14Cyr calendar provenance Lab no. source BP Ϯ SD² age at 2␴ Material Pretreatments Method ␦13C (cm) ␤ 71069 Otisville 11 640 Ϯ 60 13 850±13 420 wood acid and AMS³ Ϫ27.8 75±85 (stems) alkali ␤ 71070 Otisville 12 340 Ϯ 70 15 410±14 100 wood acid and ␤-decay Ϫ26 115±125 (stems) alkali ␤ 71071 Otisville 11 970 Ϯ 70 15 100±13 650 wood acid and ␤-decay Ϫ27.2 135±145 (stems) alkali ␤ 153148 Otisville 14 300 Ϯ 50 17 480±16 800 sediment acid and AMS Ϫ26.5 461±501 alkali ␤ 157491 Binnewater 8900 Ϯ 50 10 190±9880 sediment acid and AMS Ϫ30 1004.5±1005.5 Pond alkali AA45688 Binnewater 11 494 Ϯ 91 13 830±13 150 sediment acid and AMS Ϫ28.6 1073.5±1074.5 Pond alkali AA45687 Binnewater 11 654 Ϯ 83 14 000±13 330 sediment acid and AMS Ϫ28.5 1085.5±1086.5 Pond alkali AA45685 Binnewater 12 283 Ϯ 83 15 390±13 860 sediment acid and AMS Ϫ29 1095±1096 Pond alkali AA45686 Binnewater 12 526 Ϯ 94 15 560±14 150 sediment acid and AMS Ϫ29.6 1105±1106 Pond alkali ␤ 157492 Binnewater 13 210 Ϯ 90 16 380±15 370 sediment acid and AMS Ϫ29.4 1125±1126 Pond alkali ␤ 153148 Binnewater 22 870 Ϯ 90 NA§ sediment acid and AMS Ϫ26 1255±1256 Pond alkali ␤ 141061 Pawelski 12 180 Ϯ 60 15 285±13 845 bone (pha- collagen AMS Ϫ20.0 160±200 Farm lanx) extraction with alkali AA45690 Pawelski 11 212 Ϯ 79 13 450±12 990 sediment acid and AMS Ϫ23.5 129.5±130.5 Farm alkali AA45689 Pawelski 13 083 Ϯ 86 16 210±15 080 sediment acid and AMS Ϫ23.2 154.5±155.5 Farm alkali ␤ 135234 Hyde Park 11 480 Ϯ 60 13 805±13 165 tooth collagen AMS Ϫ22.7 enamel extraction (tusk) with alkali AA46612 Hyde Park 12 484 Ϯ 64 15 490±14 150 sediment acid and AMS Ϫ26.6 9.5±10.5 alkali AA46613 Hyde Park 13 280 Ϯ 75 16 430±15 540 sediment acid and AMS Ϫ26.5 34.5±35.5 alkali NZA12584 Temple Hill 11 000 Ϯ 80 13 173±12 661 bone collagen AMS Ϫ21.3 (NY State (scapu- extraction Museum la) and gelati- # V100) nization ² Radiocarbon years before present (where present is 1950), corrected for isotopic fractionation. ³ Accelerator mass spectrometry. § Not applicable because the date is beyond the range for calibration to calendar years.

7. Zone IVd.ÐTsuga makes a gradual recovery to Sporormiella.ÐSpores of the dung fungus Sporor- early Holocene levels by mid-zone, but is not as strong- miella spp. maintain high values, at 3±5% (Davis 1987) ly represented as in the mid-Holocene. Castanea con- throughout Zone I (Fig. 3). A steep decline begins at tinues to rise, along with Fagus and Betula, all of which the bottom of Zone II, dropping to less than 0.5% by peak at ϳ12% in the lower part of this zone, partially the middle and reaching 0% at the top of this unit. replacing Quercus, which falls to 30% for the ®rst time Above this horizon, Sporormiella is mostly absent, oc- since the late glacial. By mid-zone, Quercus has re- casionally reaching 0.5% until the late Holocene, where turned to Ͼ40%. in upper Zone IVd and lower Zone V, it climbs to 1% 8. Zone V.ÐA rapid rise in Poaceae from near zero for the ®rst time since the late Pleistocene. to 10% and the ®rst appearance of ruderal species such Charcoal.ÐThroughout Zone I, graminoid charcoal as Plantago (plantain) mark the lower boundary of this is either absent or at a very low level, while other zone. Monolete spores also show a steep rise, as does microscopic charcoal concentrations average below 2 Ambrosia (ragweed). The percentage of all tree pollen ϫ 106 ␮m2/cm 3 (see Fig. 3). Graminoid charcoal in- declined relative to upper Zone IVd. Quercus drops to creases to 5 ϫ 106 ␮m2/cm3 at the bottom of Zone II, 20% and is paralleled by decreases in Fagus, Carya, amorphous charcoal to 15 ϫ 106 ␮m2/cm3 and total Tsuga, and Pinus strobus. The only tree species to ex- charcoal reaches 20 ϫ 106 ␮m2/cm3, an order of mag- hibit an increase from Zone IVd is Betula. nitude rise overall. The subsequent pattern of charcoal August 2005 MEGAFAUNAL EXTINCTION IN NEW YORK 301

FIG.3. Sporormiella and charcoal for Binnewater Pond. Spore values are expressed as a percentage of pollen sum (upland plus aquatic). Microscopic charcoal is expressed as projected area on slide per volume of sediment. Macroscopic charcoal (Ͼ250 ␮m) is expressed as number of particles (n) per volume of sediment.

concentrations is erratic for the lower half of Zone II, tributing somewhat less than 50%. The lower boundary averaging 10 ϫ 106 ␮m2/cm3 until mid-zone, when it of Zone IVa is marked by a rise of total microscopic rises abruptly again to 75 ϫ 106 ␮m2/cm3. Amorphous charcoal to Ͼ20 ϫ 106 ␮m2/cm3, a pattern that char- charcoal rises earliest, followed closely by graminoid acterizes the remainder of Zone IV, with graminoid charcoal. By the upper part of Zone II and throughout charcoal often contributing Ͼ50% of the total. In Zone Zone III, total charcoal settles to a steady level of about V, total charcoal falls to values not recorded since Zone 10 ϫ 106 ␮m2/cm3, with the graminoid fraction con- I and lower Zone II. 302 GUY S. ROBINSON ET AL. Ecological Monographs Vol. 75, No. 3

FIG. 4. Diagram showing percentage of the total pollen and spores at each depth for major pollen and spore types for Pawelski Farm, New York, USA. The dagger symbol indicates date for the Cervalces skeleton. Cluster analysis used is constrained incremental sum of squares (CONISS). The bars to the left of the y-axis indicate the stratigraphic range from which the radiocarbon-dated material was obtained.

Pawelski Cervalces site abundant yellowish-brown, ®brous, plant material Beginning in 1996, annual ditch maintenance on the which is apparently sedges and broad-leaved plants. Pawelski farm in the Black Dirt Region of Orange The upper unit is a dark peat (Munsell color: 10YR 2/ County had uncovered several unusual bones and piec- 1), typical of the soils of New York's Black Dirt Re- es of antler. Once the source was located, systematic gion. excavation in 1999 revealed the skeleton of a male Pollen stratigraphy.ÐThe summary pollen diagram Cervalces scotti, or stag moose (Scott 1885), lying be- (Fig. 4) includes major types; zones have been estab- tween 170 cm and 200 cm below the surface. A plug lished numerically. In this instance, Poaceae and mon- of compact bone extracted from a proximal phalanx olete spores have been treated as paludal taxa. Only yielded an accelerator mass spectrometry (AMS) col- trace amounts of Poaceae pollen appear in this record lagen date of 12 180 Ϯ 60 14C yr BP (Table 3). during the late Pleistocene, but values abruptly rise to Sediment samples were taken at 10 cm intervals di- ϳ40% in Zone IVa, b, represented entirely by grains rectly from a cleaned face of the drainage ditch, above of a single thin-walled morphotype. Their importance the bone-bearing layer. Sediment below the bottom of is reduced upsection and near the surface, where mon- the ditch was sampled with a bucket auger. Subsamples olete spores approach 40%. for pollen and charcoal were taken at ϳ25-cm intervals 1. Zone I.ÐPinus is the principal taxon in this zone, for the lower part of this sequence. followed closely by Picea, and Cyperaceae, but Alnus, Sediment description.ÐDistinct boundaries delin- Quercus, and Salix are important throughout, each eate three sedimentary units. A dark blue-gray clay amounting to as much as 5%. Betula rises from a trace (Munsell color: Gley 4/N) forms the lower unit, which to above 5% at the top. Pollen tends to be sparse yet contacts a peaty clay (Munsell color: Gley 5/10Y) at adequate, except for the 298-cm sample, for which only 215 cm. The peaty clay extends to 100 cm and includes ϳ300 grains were counted in ®ve complete slides. Pres- August 2005 MEGAFAUNAL EXTINCTION IN NEW YORK 303 ervation is generally good, with grains intact but fre- change in the ®re history is marked by a charcoal con- quently crumpled, apparently from being supported in centration rise to over 25 ϫ 106 m2/cm3. Above this, a clay matrix. values then decline before rising in the middle of Zone 2. Zone II.ÐPicea rises steadily to 55%, dominating III to ϳ8 ϫ 106 ␮m2/cm3, where graminoid fragments the middle of this zone, but Pinus recovers toward the contribute a little over 50%; some macroscopic char- top. Sedges decline to low levels early in the section; coal also is present. Low levels then persist until the Betula and Quercus rise to ϳ10%. Ostrya/Carpinus lower portion of Zone IV, where microscopic charcoal (hop hornbeam and hornbeam) and monolete spores rises to 10 ϫ 106 ␮m2/cm3. Very low levels mark most display a low but signi®cant presence throughout. Salix of the Holocene record here before rising signi®cantly has a lesser presence than earlier, but keeps a steady at 20 cm from the present surface. signal to the top of Zone II. A sediment date of 13 083 Ϯ 86 14C yr BP came from near the top of the zone. Otisville The Cervalces skeleton was found within the bound- Bones of an American mastodon (Mammut ameri- aries of Zone II. canum) were uncovered in 1872 by peat diggers in a 3. Zone III.ÐPicea rises to ϳ25% mid-zone, but small bog at the foot of the eastern slope of the Shaw- drops to ϳ10%, delineating a ®nal decline. Pinus (un- angunk Mountains. The Otisville mastodon skeleton differentiated) follows the opposite pattern, decreasing now resides at the Peabody Museum in New Haven, somewhat mid-zone and peaking at the top. Pinus stro- Connecticut. Contemporary newspaper accounts and bus appears and exceeds 10% at this time, while Quer- local inquiries helped relocate the site east of the town cus declines. Alnus and Betula both rise and fall across of Otisville, New York (Table 2). The pro®le of the this zone and Tsuga appears in low numbers at the top. basin was initially traced by transecting the bog with Sediment from the bottom of Zone III dates to 11 212 a series of exploratory bucket auger cores so that an Ϯ 79 14CyrBP. appropriately deep section for excavation could be lo- 4. Zone IVa, IVb.ÐPicea disappears and Quercus cated, avoiding the area disturbed by peat mining. Sed- jumps to 60%. Tsuga rises early within the zone and iment samples and plant macrofossils were collected remains a steady presence of ϳ10±12%, while Carya, based on their stratigraphic provenance. Below 1.6 m Fagus, and Acer become a signi®cant part of the spec- depth, sediment sampling continued by using a bucket trum. Poaceae rises rapidly from trace amounts to auger down to 5.12 m, where the auger bottomed out ϳ60% of the upland and aquatic total, but is replaced in a gravelly layer, presumably consisting of glacial in the course of the upper 60 cm by monolete and drift. Osmunda fern spores, and to a lesser degree by pollen Sediment description.ÐAn AMS sediment date sets of Cephalanthus (buttonbush), a paludal shrub. Pinus a minimum time of 14 300 Ϯ 50 14C yr BP (Table 3), drops below 5% by mid-zone but recovers to exceed for onset of sedimentation at the Otisville site. A clayey 10% within 10 cm of the surface. Pinus strobus is basal gravel contacts an olive gray clay unit (Munsell present at less than 2% throughout Zone IV, with the color 5Y 4/1±4/2) at 465 cm, which becomes a bit exception of the 80 cm sample, where it barely ex- darker and more humic as it continues upsection to 320 ceeded 5%. Absent from most of Zone IV, Picea reach- cm (Munsell color 5Y 3/1±3/2), becoming a humic, es ϳ5% in the top two samples. Betula, which begins silty clay (Munsell color 5Y 4/1±4/2) up to ϳ90 cm. to decline in Zone III, now maintains a presence of From this level, peat forms the upper unit, and is dark between ϳ5% and 10% throughout Zone IV. Alnus gray, with clay from ϳ90 to 75 cm (Munsell color 2.5Y starts Zone IV at ϳ2% and does not exceed this low 3/2). Above this, the peat becomes darker up to 45 cm level until the top 30 cm, where it surpasses 5%. (Munsell color 2.5Y 4/1±4/2). It includes more woody Sporormiella.ÐThese spores are present throughout material and less clay. From 45 cm upwards, the unit Zone I, reaching 3±4% of the upland and aquatic pollen becomes darker and less woody (Munsell color 2.5 Y sum (Fig. 5). In a single sample, for which pollen was 2/0), changing to a dark gray woody peat (Munsell also sparse (see remarks on 298 cm sample, above), color 10YR 3/1) at ϳ25 cm below the surface. Sporormiella is below 1%, but otherwise is high for A surface sediment sample was taken, but much of the rest of this zone (ϳ3±4%). At the boundary with the upper 25 cm of the peat unit was disturbed by a Zone II, spore values drop to low numbers (ϳ0.5%) mass of roots and so the next sample was taken at 30 for the remainder of Zone II before disappearing at the cm, and at 10-cm intervals thereafter, to the bottom of top. With the exception of some trace amounts ob- the excavated pit. Sampling proceeded below this level served at two levels toward the center of Zone IV, by bucket auger. Sporormiella is essentially absent from the rest of the Pollen stratigraphy.ÐThe summary diagram dis- sequence. playing major pollen and spore types for the Otisville Charcoal.ÐTotal microscopic charcoal concentra- site is shown in Fig. 6. A dissimilarity coef®cient of tions in Zone I and most of Zone II scarcely exceed 2 ϳ0.6 clusters the samples into four basic zones, com- ϫ 106 ␮m2/cm3. At the top of Zone II, in the 155-cm parable to the pollen stratigraphies of Binnewater Pond sample (sediment date:13 083 Ϯ 86 14C yr BP), a and Pawelski Farm. 304 GUY S. ROBINSON ET AL. Ecological Monographs Vol. 75, No. 3

FIG.5. Sporormiella and charcoal for Pawelski Farm. Spore values are expressed as a percentage of pollen sum (upland plus aquatic). Microscopic charcoal is expressed as projected area on slide per volume of sediment. Macroscopic charcoal (Ͼ250 ␮m) is expressed as number of particles (n) per volume of sediment. The dagger symbol indicates date for the Cervalces skeleton. The bars to the left of the y-axis indicate the stratigraphic range from which the radiocarbon-dated material was obtained.

In addition to the aquatic and paludal taxa that are plant macrofossil of 11 970 Ϯ 70 14C yr BP marks the normally excluded from the upland pollen sum, Alnus top of Zone I. also is excluded from the Otisville spectra. Alder 2. Zone II.ÐPicea continues its rise, surpassing Pi- shrubs are growing abundantly on the bog surface today nus. Betula rises and peaks before mid-zone, and Alnus and polyads of more than 60 Alnus grains are found in declines steadily, as noted at the Binnewater and Paw- fossil samples. Monolete spores also are excluded for elski sites. Quercus and Poaceae form a steady ϳ5± similar reasons. 8%. Salix has a similar value, but is somewhat higher 1. Zone I.ÐPinus and Picea, in that order, dominate at mid-zone. Abies and Pinus strobus appear. Artemisia these late-glacial pollen spectra. Toward the top, Picea and Thalictrum drop to lower levels, but Poaceae and rises at the expense of Pinus. Broadleaf taxa include Asteraceae (high echinate types) have a presence com- Betula, which exceeds ϳ15% in the upper part of the parable to that of Zone I. zone. Quercus, Salix, and Poaceae each attain 8±10%. 3. Zone III.ÐPinus now dominates the assemblage, Cyperaceae often exceeds 20%. Thalictrum reaches followed by Picea, which drops out abruptly at the top 5%, forming part of a tundra community along with of this zone. Alnus reaches almost 10% and Betula, Artemisia and other Asteraceae. An AMS date on a 15%. Tsuga appears for the ®rst time and Quercus re- August 2005 MEGAFAUNAL EXTINCTION IN NEW YORK 305

FIG. 6. Diagram showing percentage of the total pollen and spores at each depth for major pollen and spore types for Otisville, New York, USA. Dagger symbols indicate plant macrofossil dates. Cluster analysis used is constrained incremental sum of squares (CONISS). The bars to the left of the y-axis indicate the stratigraphic range from which the radiocarbon- dated material was obtained. mains at ϳ8%. A plant macrofossil AMS dated at sample above this horizon except for the surface, where 11 640 Ϯ 60 14C yr BP originated from the lower sam- it reappears in low numbers. Overall, Sporormiella val- ple. As in Binnewater Pond and Pawelski Farm, the ues of the Pleistocene record at Otisville are not as last appearance of Picea may be seen as de®ning the high as at Pawelski Farm or Binnewater Pond, but nev- top of this zone. ertheless are comparable with those of modern sedi- 4. Zone IV.ÐPinus continues to dominate at 40%, ments in overstocked areas (Davis 1987, Davis and and at the bottom of this zone Picea is absent. Pinus Shafer 2002). In contrast to other sites, Sporormiella strobus, although registering a trace presence earlier, maintains a continuous although declining presence at exceeds 10% in the 60-cm sample that forms the bottom this site through Zone III and ®nally drops out at the of the zone. Above this point Pinus falls to 15% and same stratum as Picea. P. strobus below 2%. Quercus climbs rapidly to 40%, Charcoal.ÐOtisville contains background levels of Betula drops below 5%, and Tsuga rises to 15%. Carya amorphous charcoal that are many times higher than and Castanea appear midzone and each exceed 5%; elsewhere (Fig. 7). From the beginning of sedimenta- monolete spores become abundant. tion, at the bottom of Zone I, charcoal concentration Sporormiella.ÐThe fungal spores associated with is ϳ50 ϫ 106 ␮m2/cm3, and even though this level dung have a signi®cant presence of ϳ2±3% throughout declines to about 20 ϫ 106 ␮m2/cm3 by the top of Zone Zone I, show somewhat lower values of ϳ1±2% I, it rises again to ϳ35 ϫ 106 ␮m2/cm3 in Zone II, and through Zone II, and fall to their lowest encountered averages only slightly less for the remainder of the Late level of ϳ0.2% near the top. They subsequently recover Pleistocene. Macroscopic charcoal appears ®rst in Zone slightly before disappearing from the record at the top III (Younger Dryas) and then in the surface sample. of Zone III. Sporormiella then is absent from every Microscopic charcoal concentrations rise much later 306 GUY S. ROBINSON ET AL. Ecological Monographs Vol. 75, No. 3

FIG.7. Sporormiella and charcoal for Otisville. Spore values are expressed as a percentage of pollen sum (upland plus aquatic). Microscopic charcoal is expressed as projected area on slide per volume of sediment. Macroscopic charcoal (Ͼ250 ␮m) is expressed as number of particles (n) per volume of sediment. Dagger symbols indicate plant macrofossil dates. The bars to the left of the y-axis indicate the stratigraphic range from which the radiocarbon-dated material was obtained. than at all other sites, beginning at 50 cm from the until the following year when arrangements were made surface and coinciding with the Quercus peak in Zone to excavate below the water table, permitting more con- IV. trolled removal of matrix sediments (D. A. Burney and G. S. Robinson, unpublished manuscript), which re- Hyde Park vealed a nearly complete skeleton of an adult male During late summer of 1999, a backhoe deepened an Mammut americanum. The thoracic spinal vertebrae, arti®cial pond in a wetland area of suburban Hyde Park, pelvis, and sacrum were substantially articulated and Dutchess County, New York, and encountered the limb a fully articulated foot was positioned plantar side bones of a probiscidian. A fragment of tusk recovered down in the sediments. from the spoil heap at the site yielded an AMS collagen Inspection of a cleaned vertical face of the pit date of 11 480 Ϯ 60 14C yr BP (Table 3). Work stopped showed that while the backhoe may have removed August 2005 MEGAFAUNAL EXTINCTION IN NEW YORK 307

FIG. 8. Diagram showing percentage of the total pollen and spores at each depth for major pollen and spore types for Hyde Park, New York, USA. The dagger symbols indicate plant macrofossil dates. Cluster analysis used is constrained incremental sum of squares (CONISS). The bars to the left of the y-axis indicate the stratigraphic range from which the radiocarbon-dated material was obtained. much of the upper sedimentary units, the underlying consisting of clasts up to 1 cm diameter. At 55 cm, on stratigraphy was intact. Samples for microfossil work a clear boundary, the humic clay unit meets a grayish were removed directly from the face, at 5-cm intervals. brown (Munsell color: 2.5Y5/2) humic silty clay that A parallel set of large volume samples at 10 cm inter- contains a calcareous molluscan gravel. At about 30 vals were taken for macrofossil analysis. Sampling cm, there is a laterally uneven contact with a dark gray- from 100 cm below the sediment surface (below the brown silty clay (2.5Y4/2), which includes wood, ¯oor of the excavated pit) was continued at 5-cm in- grass, and sedge ®ber; broken snails; and other mollusk tervals using a 7.5-cm bucket auger. Fist-sized cobbles shells. Another uneven contact occurs at about 10 cm, were encountered at 175 cm, requiring the use of a characterized as a coarse detrital peat, which bears posthole digger to remove material down to 190 cm, some clay, but little or no sand. A sediment date of at which point the sediment became too indurated for 12 484 Ϯ 64 14C yr BP originated from the bottom of further removal. the peat member, but a similarly provenanced Abies Sediment description.ÐThe basal sandy gravel in- cone scale yielded an AMS date of 11 230 50 14Cyr cludes cobbles of subrounded gray sandstone of 6 cm BP. A tracing of this peat unit laterally shows that much average diameter. Smaller stones, up to 4 cm diameter, of its upper part was removed by a backhoe bucket. are present in the unit from 175 cm to 160 cm, where Pollen stratigraphy.ÐThe entirely late Pleistocene a uniform dark bluish gray clay (5B 4/1) is contacted, pollen stratigraphy of this site clearly is divided into also uniform in appearance, up to 140 cm where it two pollen zones (Fig. 8). meets a dark gray clay (5Y 4/1). At 100 cm, the clay 1. Zone I.ÐPinus dominates throughout, and makes becomes more humic, containing some plant ®bers and up as much as 60% of the lower part of this zone, where degraded molluscan gravel. It is interrupted at 80 cm it is followed by Picea (19%), Alnus (8%), Quercus by a lens of coarse sand and angular schistose gravel (5%), and Salix (2%). Together with the less abundant 308 GUY S. ROBINSON ET AL. Ecological Monographs Vol. 75, No. 3

Poaceae and Artemisia, these latter taxa begin to con- concentration at 45 cm is 2.6 ϫ 106 ␮m2/cm3, subse- stitute a larger share in the upper part of the zone. quently rising to 28 ϫ 106 ␮m2/cm3 at 35 cm, with Trilete spores are present earlier; monolete spores and graminoid and amorphous types at approximately sub- Lycopodium sporadically occur throughout the pro®le. equal contributions. Charcoal values decline above Cyperaceae rise steadily from 10% to 25%. The pres- this, but remain at ®ve to 10 times the average found ence of Dryas is entirely within Zone I; it comprises in Zone I. Macroscopic charcoal ®rst appears at the ϳ3% of the lowest sample and trace amounts are found bottom of Zone II, then rises at 35 cm. throughout the zone. 2. Zone II.ÐPicea rises abruptly to 55% and ex- Temple Hill mastodon ceeds 70% at mid-zone, declining to about 40% to- One of many mastodons found in Orange County, wards the top. Pinus spp. is less than 20% of the sum, the Temple Hill specimen was excavated in 1921, after although the occurrence of Pinus strobus (haploxylon) it was encountered during trenching in mucklands is a consistent, if low, presence from the mid-zone on about 4 miles (6.4 km) west of Newburgh, New York up. Abies appears slightly earlier, and exceeds 5% at (Hartnagel and Bishop 1922). The largely complete the top. Quercus rises steadily to about 18%. Betula skeleton is now at the State Museum of New York in and Carpinus/Ostrya rise and maintain a level of ϳ5% Albany. A plug of compact bone removed from the and 2%, respectively. Alnus and Salix decline early on subscapular fossa yielded an AMS age of 11 000 Ϯ 80 but do not disappear. High echinate Asteraceae and 14C yr BP (Table 3). Poaceae are present throughout the interval, as in Zone I; Artemisia declines but remains in the pro®le. Mon- DISCUSSION olete spores are present throughout, averaging less than Biostratigraphy 5%. Cyperaceae is less than 10% at the bottom of Zone II but increases to ϳ15% upsection. Myriophyllum (wa- The sedimentary histories of each site display key ter milfoil) appears at the bottom, rises ca. 4%, declines features of the late-glacial pollen stratigraphy of south- ern New England (Deevey 1939, 1943, Leopold 1956, at midzone, but is still present at the top. Davis 1969). This correspondence is most complete for The sediment date of 13 280 Ϯ 75 14CyrBPwas Binnewater Pond, which singly received uninterrupted taken at 35 cm depth, in the lower half of this zone, sedimentation during the late Holocene. Nevertheless, but an AMS age on Picea needles located ϳ10 cm all sites incorporate a microfossil stratigraphy that below this was 12 300 Ϯ 40 14C yr BP (N. G. Miller, brackets the extinction window. unpublished manuscript). The following observations Zone I at Binnewater Pond is equivalent to pollen suggest that Zone III is missing, along with a portion zone A1 of the southern New England sequence. Sim- of Zone II and much of the peat unit of the sedimentary ilarly, Zone II corresponds to zones A2±A3, re¯ecting strata. Although Picea shows a slight decline near the the late-glacial rapid warming of the BoÈlling/AlleroÈd top the Zone II, Quercus has risen to 18% and displays Interval (Peteet 1992); Zone III is equivalent to A4, no sign of decline; Tsuga remains absent. There is no now recognized as a vegetational response to the Youn- pronounced rise in Betula as in the other described sites ger Dryas cooling event in the northeastern United that would be expected to mark Zone III. Alnus, which States (Leopold 1956, Peteet et al. 1990, Peteet 1992). has fallen from its strong presence in Zone I, shows In particular, Zone III pollen spectra display a char- no recovery by the top of the sequence. acteristic rise then fall of Alnus, that marks the begin- A particularly rich record of plant macrofossils re- ning and end of the Younger Dryas across regions from ported elsewhere (N. G. Miller, unpublished manu- New Jersey to the Atlantic Provinces of Canada (Mayle script), is in close agreement with the pollen stratig- et al. 1993). Zone IVa matches the C1 pollen zone, a raphy described here. response to the onset of Holocene warming. Sporormiella.ÐIn common with the other sites, at Other key points of correspondence that occur in the Hyde Park, Sporormiella has a strong presence remainder of the postglacial sequence include the Tsu- throughout Zone I, even exceeding 6% (Fig. 9) at some ga decline, an event recorded in pollen diagrams from horizons. At the bottom of Zone II, the spores begin a sites throughout the range of Tsuga canadensis. Dating steady decline, and while never quite reaching zero, to ca. 4800 14C yr BP, this ``hemlock decline'' has been values are very low by the top of Zone II, as at Bin- attributed to a forest pathogen outbreak, (Davis 1981, newater Pond and Pawelski Farm. Following the initial Allison et al. 1986), and is a useful stratigraphic marker decline in the middle of Zone II, spores brie¯y return for correlating sequences across eastern North Amer- to 6% in a single sample (25 cm) then drop to a less ica. than 0.5% average for the remainder of the sequence. The Ambrosia horizon, marking the boundary of Charcoal.ÐThroughout Zone I (Fig. 9), all charcoal Zone V, is prevalent throughout the Northeast and much is below 1 ϫ 106 ␮m2/cm3 and graminoid charcoal is of the Midwest and is understood to be a cultural rather at trace levels. In the lower part of Zone II, an order than climatic signal for the late Holocene pollen record. of magnitude rise occurs. Total microscopic charcoal Together with other palynological markers, the rise of August 2005 MEGAFAUNAL EXTINCTION IN NEW YORK 309

FIG.9. Sporormiella and charcoal for Hyde Park. Spore values are expressed as a percentage of pollen sum (upland plus aquatic). Microscopic charcoal is expressed as projected area on slide per volume of sediment. Macroscopic charcoal (Ͼ250 ␮m) is expressed as number of particles (n) per volume of sediment. Daggers indicate plant macrofossil dates. The bars to the left of the y-axis indicate the stratigraphic range from which the radiocarbon-dated material was obtained.

Ambrosia re¯ects land-use changes associated with Eu- taining units. Each of these samples had low carbonate ropean settlement about 300 years ago in the Northeast content (Robinson 2003), and they were rejected for (Miller 1973, Russell 1980, Peteet 1992). the following reasons. The date of 22 850 Ϯ 90 14Cyr Most New England pollen stratigraphies include a BP is not consistent with a location signi®cantly north ``B'' or Boreal Zone typically de®ned by a ®nal Pinus of the glacial margin until at least ca. 17 000 14CyrBP peak occurring just above the Younger Dryas. In stud- (Connally and Sirkin 1970). The date of 13 210 Ϯ 90 ies where P. strobus or haploxyon pine pollen is dis- 14C yr BP is from a pollen stratigraphic horizon cor- played separately, the ``B- Zone'' Pinus peak is formed relative with the earliest A2±A3 pollen zone. Other primarily if not exclusively by P. strobus, or white pine studies from the Wallkill and lower Hudson valley re- (Davis 1969). An equivalent of this zone notably is gion put this horizon at ca. 12 600 14C yr BP (Connally absent from the Binnewater Pond sequence, although and Sirkin 1970, Peteet et al. 1993, Maenza-Gmelch it is clearly re¯ected in two of the other sites of this 1997a, b). While hard-water error is unlikely in these study. cases, glacial clay may have introduced ``in®nite-age'' Although all the sediment dates from the Binnewater carbon into the organic fraction (Sutherland 1980) and core are internally consistent, two are from clay-con- thus invalidated the results. 310 GUY S. ROBINSON ET AL. Ecological Monographs Vol. 75, No. 3

Sporormiella declined at Binnewater Pond at the bot- sediment has been burned off and is regularly de¯ated tom of Zone II, suggesting an abrupt drop in the density by winds, exposing layers laid down at least 5000 years of large herbivores around this site during the early ago. Because of arti®cial drainage, the surface is dry stage of the BoÈlling/AlleroÈd late-glacial warming. Near for much of the year, which explains why it does not the boundary of Zones IV and V, presumably at around sequester a modern pollen rain. the time of European settlement, Sporormiella values The sediment date marking the Zone II charcoal peak surpass any samples since the late Pleistocene. There is not consistent with the pollen stratigraphy, which is are two maxima: the ®rst possibly caused by relaxed clearly younger. However, the Cervalces skeleton was predation pressure on game animals when native Amer- recovered from 200 cm depth and AMS dated to 12 180 ican populations were decimated by epidemics follow- Ϯ 60 14C yr BP. Although it is possible that the bones ing ®rst European contact. The second peak may mark sank into the sediments, the collagen date is consistent the subsequent introduction of domestic ungulates (cf. with the pollen spectrum of its matrix. Accordingly, Davis 1975). the date marks the interval between Sporormiella de- Immediately preceding the Late Pleistocene drop in cline at the Zone I±Zone II boundary, and the charcoal Sporormiella, there is an order-of-magnitude rise in peak that occurs ϳ60 cm above it. both graminoid and other microscopic charcoal. This The pollen spectrum of Otisville's Zone II correlates rise occurs at the boundary of Zones I and II, and may with the BoÈlling/AlleroÈd warming, an interpretation be the ®rst sign of human activity in the region (the supported by the radiometric date of 12 340 Ϯ 70 14C second being Sporormiella decline). The largest in- yr BP on a plant macrofossil. However, because of crease in ®re frequency appears later, about 60 cm high- radiocarbon anomalies that characterize this time pe- er, above the middle of Zone II in the Binnewater se- riod (Fiedel 1999), the stratigraphic reversal of this date quence, at 12 283 Ϯ 83 14C BP, on AMS dated sedi- with the later one from Zone I is more apparent than ments, and may be a result of litter accumulation fol- real. When calibrated at 2 sigma the two dates in ques- lowing the depletion of large herbivores. By Zone V, tion overlap by a thousand years (Table 3). Otisville's the charcoal data indicate that ®re is now reduced in Zone III is represented by two samples showing a peak importance to a level not seen for ca. 13 millennia. in Betula and Alnus, and establishes the Younger Dryas At Pawelski Farm, the low concentration of pollen chronozone. in the lower unit may be due to rapid deposition of Otisville's pollen stratigraphy shows a clear Pinus clay from glacial melt. The pollen pro®le of Zone III strobus spike, equivalent to the boreal or ``B Zone'' of is indicative of the Younger Dryas chronozone, partic- the New England pollen stratigraphy. It appears at 60 ularly the unprecedented rise and then fall in Alnus cm depth and cluster analysis (dendrogram, Fig. 6) rec- pollen spanning the unit. The late glacial and early ognizes it as a zone in its own right from a dissimilarity Holocene pollen stratigraphy correlates well with that coef®cient of ϳ0.9. However, at a larger dissimilarity of Binnewater Pond, although inspection of Fig. 4 coef®cient it clusters with Zone IV, to which we have shows that, based on a dissimilarity coef®cient of ϳ0.8 assigned it here, with a broken line indicating its upper chosen in this case to establish pollen stratigraphic limit. zones, the single sample at 100 cm depth would also As elsewhere, the early Holocene warming is marked qualify as a zone in its own right. By correlation with by a signi®cant rise in Quercus, but sedimentation has New England pollen stratigraphy, (Davis 1969), this is slowed above this, so the later Holocene is poorly rep- the ``B-Zone'' that is apparently missing or obscured resented. However, all taxa present at 30 cm also appear by other taxa in the Binnewater Pond pollen pro®le. in the surface sample in roughly the same percentages, As in New England, it is de®ned here by a rapid rise the principal exception being Ambrosia, constituting of Pinus strobus to ϳ45%. This ``boreal'' zone is un- ϳ8% at the surface but barely present at 30 cm. This derstood to postdate the Younger Dryas chronozone provides some assurance that root disturbance has not (Peteet et al. 1993), and is demarcated here by a broken introduced late Holocene material into the lower por- line toward the top of Zone III in Fig. 4 and Fig. 5. tion of Zone IV. The abundance of Ambrosia and Cas- The striking rise of Poaceae pollen and later of mon- tanea at the surface suggest that it is late Holocene and olete spores is accompanied by aquatics suggesting lo- likely of post-European contact time. Thus, it might cal changes in the wetland plant community. Quercus have been assigned to Zone V, but as a single sample dominance signals the onset of Holocene warming, fol- that did not cluster by the chosen dissimilarity coef- lowed by other thermophilous species. Today, the top ®cient, we have grouped it with Zone IV. ϳ30 cm of the peat unit is disk plowed and planted Otisville's charcoal record indicates that ®re was an with onions, and yet the uppermost pollen spectra have important element of the local ecology by the time the remained well strati®ed, suggesting that such distur- earliest sediments began to accumulate. When micro- bance does not signi®cantly mix the sediments verti- scopic charcoal concentrations rise, they reach levels cally. Furthermore, the pollen spectrum at the surface not seen at other sites: 210 and 260 ϫ 106 ␮m2/cm3 for is not modern: the mid-Holocene Tsuga decline and amorphous and graminoid types respectively. As a the Ambrosia horizon are missing. Much of the surface small sedimentary basin, this site would be expected August 2005 MEGAFAUNAL EXTINCTION IN NEW YORK 311 to receive a strong local signal in its charcoal record. horizon, an Abies cone scale is AMS dated only slightly Nevertheless, that record is not one of sporadic and younger than the mastodon itself (Fig. 9). Although occasional ®res that might be over-represented, but of this could suggest a brief return of megafauna to this frequent natural ®res that are a consistent feature of area, a simpler explanation is that the brief in¯ux of the local ecology. The ®rst macroscopic charcoal frag- Sporormiella migrated from the gut of the mastodon ments appear somewhat earlier, in upper Zone III, in- that collapsed and died at about this time only ca. 1 m dicating burning close to the site (Patterson et al. 1987). away. In a small sedimentary basin, a high density of Sporormiella remains moderately high (over 1%) dur- spores may have had little opportunity to extensively ing most of late Pleistocene time, up into lower Zone disperse. III, before falling to zero at the same time as Picea.A reappearance of the spores and Ambrosia at the surface Hypotheses for extinction suggests introduced livestock and European agricul- If climate change was the suf®cient cause of extinc- ture. Dairy cattle (of the Mitchell farm) occupied the tion (H1 in Table 1; see Appendix) we should expect site at least from the 19th century (H. R. Decker, per- Sporormiella to decline at all sites simultaneously. sonal communication). Even at this spatial scale, spore decline is not simul- Zone II at Hyde Park encompasses the spruce max- taneous, whether measured by pollen chronostratigra- imum or A2±A3 zones of the regional pollen stratig- phy or by absolute dates. Of the two sites that have raphy (Peteet et al. 1993, Davis 1969, Maenza-Gmelch plant macrofossil dates, Hyde Park and Otisville, one 1997a, b) as well as of northern New Jersey (Peteet et shows a Sporormiella decline at least a millennium al. 1990), re¯ecting the BoÈlling/AlleroÈd warming (Pe- earlier than the other. Most sites record this event ear- teet 1992). However, this is not followed by an Alnus lier or later in pollen Zone II, but at Otisville, Spo- maximum, indicating that the Younger Dryas is not rormiella persists through Zone III, only to disappear represented. Pollen and macrofossils of rooted aquatic at the same time as Picea pollen. There is no suggestion taxa such as Myriophyllum suggest the formation of a at this stage of the formation of a uniform or zonal shallow pond (N. G. Miller, unpublished manuscript). vegetation pattern (Guthrie 1984). Such a development Microscopic charcoal particles of Hyde Park Zone I occurs yet in the future, at ca. 9000 14C yr BP, when are of small size classes, mostly between 50 to 500 Quercus rapidly dominates regional pollen spectra, ␮m2 (Robinson 2003). This continues through lower suggesting the formation of closed-canopy forest at the Zone II, suggesting a time of not only less frequent beginning of the Holocene ®re, but that charcoal originated from more distant Sporormiella data from three of the four sites indi- sources (Clark 1988). Larger size classes and macro- cate that large herbivores were in decline at the very scopic charcoal of at least 250 ␮m (Fig. 5) take a greater time that Picea (spruce) was either rising or already share at 50 cm depth, a pattern indicating local ®res dominating regional pollen spectra. This tends to con-

(Clark 1988). However, concentrations remain low tradict H2, which holds that the American mastodon overall, and consequently by this time ®re was not a became so dependent on spruce that it died out as hard- frequent occurrence. A macrofossil date suggests that wood forests expanded. after 12 300 Ϯ 40 14C yr BP, ®re abruptly became fre- At least a millennium elapsed between the earliest quent; an increase in average particle sizes also sug- local megafaunal collapse (Sporormiella decline) and gests that both local and regional ®re contributed to dates for extinct megafauna reported here (Table 3). the rise in charcoal concentration. Graminoid (grass Recently obtained AMS collagen dates from New and sedge) charcoal accounts for most of the largest York's Fingerlake region are 11 630 Ϯ 80 14CyrBP size classes. Charcoal rise at Hyde Park stratigraphi- and 10 840 Ϯ 60 14C yr BP for mastodon, and 10 890 cally follows closely upon Sporormiella spore decline Ϯ 50 14C yr BP for mammoth (P. Nester, personal com- in the same manner as observed in the records of Bin- munication). Even if megaherbivores declined rapidly newater Pond, Pawelski Farm, and Otisville. at individual sites, a rapid wave did not sweep through Plant macrofossil dates suggest that Sporormiella de- the entire region. These observations weaken all hy- cline at Hyde Park cannot be much earlier than 12 300 potheses predicting a rapid extirpation of megafauna 14 C yr BP (Fig. 9 and N. G. Miller, unpublished man- on the regional scale, speci®cally H3,H4,H6,H9,H10, uscript). Similar dates from southern New England and H12 of Table 1. A further dif®culty for H3 and H9 constrain the corresponding pollen horizon at ca. is that megafaunal populations appear to have gone into 12 590 Ϯ 430 14C yr BP (Peteet et al. 1992). The ter- steep decline well before the Younger Dryas climate restrial record from northwestern Europe places the reversal. Even if large animals were eliminated from onset of the BoÈlling at ca. 13 000 14C yr BP (Watts wide areas, pockets apparently remained, suggesting a

1980). The signi®cance of an apparently later Sporor- mosaic pattern of extirpation, as expected from H5 and miella decline at Otisville is addressed below. H13.

An anomaly in the Sporormiella record at Hyde Park The overkill model of H8 calls for a tripartite pulsed occurs at 20 cm, where, following initial decline, spore extinction over a period of ca. 1640 years, but the gen- values brie¯y return to a maximum. Just above this erally unimodal drop in Sporormiella values presented 312 GUY S. ROBINSON ET AL. Ecological Monographs Vol. 75, No. 3 here re¯ects only one of the two major pulses predicted. cores from across the island display a diachronous Spo- The most abundant late occurrences in the northeast rormiella decline, commencing in the arid southwest, region are mastodon, a member of the megaherbivore ca. 2000 years ago. At each site, spore decline is fol- guild, predicted by this model to disappear in the mid- lowed by a sharp rise in stratigraphic charcoal, suc- dle round. ceeded by the last occurrences of large herbivore taxa. The ``second order predation'' overkill model cannot This occurred in a background of climatic aridi®cation account for the repeated conjunction of a terminal Spo- that was already well underway prior to the advent of rormiella decline followed by a charcoal rise. H7 pre- humans. dicts that initial overkill of megaherbivores (and pos- The Madagascar records also assist resolution of un- sibly carnivores) would be followed by population out- certainty regarding the signi®cance of the Zone II char- breaks of smaller herbivores that then rapidly exhaust coal rise at Hyde Park, Pawelski, and Binnewater, when available vegetation. If it was a more regional phe- spruce is either dominant or on the decline. The spruce nomenon, the spore anomaly at Hyde Park might sup- pollen rise, found throughout formerly glaciated parts port H7 and H8. of the Northeast is considered to mark the transition None of the single-causal hypotheses are of much from periglacial tundra to open conifer woodland help in explaining why a drop in Sporormiella is fol- (Watts 1983). This development may seem suf®cient lowed by a rise in stratigraphic charcoal concentrations. for ®re to become an important factor on the local

However, H11 proposes that large herbivores roaming landscape. In Madagascar, however, a very different continental landscapes would have kept fuel loads low set of environmental changes were present, yet a char- by consuming vast quantities of biomass (SchuÈle 1990). coal rise appears linked to spore decline near the time If such animals were removed by some novel factor of human arrival. such as human activities or introduced disease, frequent Stratigraphic charcoal records from the Hudson and intense ®res might result. Highlands that overlook the eastern Wallkill Valley also

Synergy (H13) incorporates the above view of her- suggest that environmental changes alone do not ac- bivory and ®re but extends the hypothesis to a range count for all the available data. These upland sites of human activities. These activities include hunting would be more prone to lightning strikes, but show a and landscape ®res that constitute an extinction process later onset of ®re than most of the lowland areas of that initially would be expected to be relatively slow, this study. It took until the early Holocene for ®re to but later cumulative and may then be compounded by play a signi®cant role at Sutherland Pond, at 380 m ``natural'' environmental shifts. Binnewater Pond, the elevation (Maenza-Gmelch 1997a). By contrast, highest resolution sequence, records a brief, 10-fold Spruce Pond at 223 m elevation, incipiently records rise in charcoal that immediately precedes Sporor- frequent ®re in Younger Dryas time (Maenza-Gmelch miella decline. This possibly is the ®rst sign of earliest 1997b). Together with Otisville, these records begin to human settlers locally, who then overexploited avail- outline a regional ®re history that was not tracking able game, undercutting an established herbivory re- climate changes in any simple fashion, but rather sug- gime, and completing a ®rst stage of landscape trans- gest a mosaic pattern of change that might be elucidated formation. In the second phase, according to charcoal with further Sporormiella studies. data, accumulating fuel loads develop a potential for Dates on organic material associated with the earliest more pervasive and frequent ®res that appear later. A ¯uted point tradition, ca. 11 500 14C yr BP, are still the second well-known prehistoric activity, the deliberate most widely accepted interval for human arrival in the use of ®re to clear underbrush and encourage a renewed Americas south of the ice sheets (Haynes 1969, Fiedel ¯ush of growth attractive to game (Patterson and Sas- 1999). Generally accepted dates for the settlement of saman 1988), could have ampli®ed the earlier devel- eastern North America remain no earlier than ca. 11 000 opments. Although major environmental shifts mark 14C yr BP (Dincauze 1993). late-glacial time in the Northeast, a subsequent event The Dutchess Quarry Caves are located in the Wall- could have a multiplied effect for a system already kill Valley ϳ8 km northeast of the Pawelski farm and under a sustained assault. We propose that this is why ϳ13 km southeast of Binnewater Pond. Formed in a terminal dates of most extinct taxa cluster around the dolomitic limestone outcrop, the caves have yielded onset of the Younger Dryas chronozone, apparently a Paleo-Indian projectile points and remains of the ex- millennium or more after people have entered the re- tinct giant beaver, ¯at-headed peccary, caribou, and gion. Paleolithic activities may be the underlying and other extant vertebrate taxa (Funk 1976, Funk and suf®cient cause of the observed landscape transfor- Steadman 1994). Several AMS 14C ages range from mation, but such effects would be compounded by the 13 840 Ϯ 80 to 11 670 Ϯ 70 14C yr BP, on puri®ed inevitable climate changes. collagen from caribou bones recovered from or above It is useful to compare these New York records to stratigraphic units bearing ¯uted projectile points Late Holocene deposits in Madagascar, which track en- (Steadman et al. 1997). However, there is no clear as- vironmental and other changes leading up to and fol- sociation between the cultural material and extinct lowing human arrival (Burney et al. 2003). A series of megafaunal species. Questions remain about the integ- August 2005 MEGAFAUNAL EXTINCTION IN NEW YORK 313 rity of the Dutchess Quarry Cave stratigraphy. Al- here as primary causes. Once human activities were though the proxy data of this study indicates otherwise, brought to bear on the system, the effects of ``natural'' no direct evidence has emerged to place settlement in stresses could then have been ampli®ed (Burney the region before 11 000 14CyrBP. 1993a, b, 1999). The spore data in particular, suggest an initial blitzkrieg (Martin 1984) as a crippling but SUMMARY AND CONCLUSIONS not fatal blow. Human use of ®re may have ampli®ed Stratigraphic pollen, spore, and microscopic char- the ecological response to a severely reduced herbiv- coal records from four sites in southeastern New York ory. Climate change appears to have compounded these put a regional focus on the mass extinction of large effects, culminating in the disappearance of all the larg- mammals in North America. The proxy data from four est members of the North American mammalian fauna. localities show a substantial agreement regarding ACKNOWLEDGMENTS changes in herbivory and ®re regimes that unfolded Costs of radiocarbon dating were underwritten by the New during the late Pleistocene. A region-wide drop in Spo- York State Museum at Albany and the NSF Arizona AMS rormiella values during the late glacial is associated Facility of University of Arizona. In this regard, we thank everywhere with a subsequent rise in stratigraphic Tim Jull and Mitzi de Martino. Other costs were borne by charcoal. NSF grants to D. A. B.'s Laboratory at Fordham University, the Ludwig and Paula Altman Fund, the Fordham Graduate Correlation of these data with pollen stratigraphy Student Association and an Alumni Dissertation Fellowship implies that human alteration of the landscape and awarded by the Graduate School of Arts and Sciences of megafaunal collapse began signi®cantly before the Fordham University. Paul Martin actively encouraged the monumental changes in vegetation that mark the Ho- work and commented on early drafts. Thanks also go to Dorothy Peteet, George Dale, Craig Frank, James Lewis, locene boundary. Microfossil stratigraphy is supported and Berish Rubin. Robin Andersen, Gordon Connally, Tom by the radiocarbon chronology presented here to show Daniels, and Mary Kay O'Rourke have each made valuable that regional ®re and herbivory regimes were trans- comments. The manuscript bene®ted from suggestions by formed in advance of the Younger Dryas climatic re- Owen Davis and two anonymous reviewers. Access to ®eld versal. sites or museum specimens was made possible with the as- sistance of Ray Decker, Brian Pawelski, Vivian Kagan, Bone collagen dates on extinct megafauna as late as Larry and Sherry Lozier, Gay Malin, Don Cadwell, and the 11 000 Ϯ 80 14C yr BP are seen as the terminal stage late Bob Funk. The of®cers and staff of the Paleontological of a process in which ecological collapse followed hu- Research Institution, particularly Peter Nester, Jim Sherpa, man arrival, leaving severely reduced populations of Robert Ross, and Warren Allmon were responsible for man- aging the excavation at Hyde Park. Norton Miller kindly megafauna that survived to the end of the Pleistocene. granted permission to cite plant macrofossil dates before The preceding observations may be used to evaluate publication. the proposed causes of megafaunal extinction in North LITERATURE CITED America (Table 1). Hypotheses that call for an extinc- Alley, R. B., J. Marotzke, W. D. Nordhaus, J. T. 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APPENDIX A review of the proposed causes of the Late Pleistocene extinctions in North America is presented in ESA's Electronic Data Archive: Ecological Archives M075-011-A1.