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Robinson, G.S., L.P. Burney, and D.A. Burney, 2005. Landscape

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Robinson, G.S., L.P. Burney, and D.A. Burney, 2005. Landscape Ecological Monographs, 75(3), 2005, pp. 295±315 q 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.
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