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Test of Martints Overkill Hypothesis Using Radiocarbon Dates on Extinct Megafauna

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Test of Martints Overkill Hypothesis Using Radiocarbon Dates on Extinct Megafauna Test of Martin’s overkill hypothesis using radiocarbon SPECIAL FEATURE dates on extinct megafauna Todd A. Surovella,1, Spencer R. Peltona, Richard Anderson-Sprecherb, and Adam D. Myersc aDepartment of Anthropology, University of Wyoming, Laramie, WY 82071; bDepartment of Statistics, University of Wyoming, Laramie, WY 82071; and cDepartment of Physics and Astronomy, University of Wyoming, Laramie, WY 82071 Edited by Yadvinder Malhi, Oxford University, Oxford, United Kingdom, and accepted by the Editorial Board August 5, 2015 (received for review February 26, 2015) Following Martin [Martin PS (1973) Science 179:969–974], we pro- From this premise, we make two complementary arguments pose the hypothesis that the timing of human arrival to the New pertaining to New World colonization. First, we expect initial World can be assessed by examining the ecological impacts of a small megafaunal declines for each region to correspond temporally population of people on extinct Pleistocene megafauna. To that end, with the first evidence of human presence. Second, we expect that we compiled lists of direct radiocarbon dates on paleontological the timing of megafaunal declines should be geographically pat- specimens of extinct genera from North and South America with terned according to Martin’s (1) model in which the founding the expectation that the initial decline of extinct megafauna should population moved through EB and then south into the CUSA and correspond in time with the initial evidence for human colonization SA. When Martin published his classic work, radiocarbon cali- and that those declines should occur first in eastern Beringia, next in bration was not possible, so by necessity, he worked within the the contiguous United States, and last in South America. Analyses of radiocarbon time scale. For the CUSA, Martin estimated a mean spacings and frequency distributions of radiocarbon dates for each colonization age of ca. 11,200 14C y BP, or ca. 13,100 BP, and for region support the idea that the extinction event first commenced in SA, ca. 10,700 14C y BP, or ca. 12,700 BP. Martin did not specify a Beringia, roughly 13,300–15,000 BP. For the United States and South human arrival date for EB, except to suggest his model required America, extinctions commenced considerably later but were closely that “the time of human entry into Alaska need be no older than spaced in time. For the contiguous United States, extinction began at 11,700 [radiocarbon] years ago,” or roughly 13,600 BP. BIOLOGY ca. 12,900–13,200 BP, and at ca. 12,600–13,900 BP in South America. Importantly, the expectation of a north to south spatiotem- POPULATION For areas south of Beringia, these estimates correspond well with the poral extinction trend across the Western Hemisphere should be first significant evidence for human presence and are consistent with largely unique to the overkill hypothesis. There is no single cli- the predictions of the overkill hypothesis. matic (8, 9) or catastrophic (10) extinction hypothesis that shares this prediction. Therefore, this kind of analysis is not only ca- Pleistocene extinctions | overkill | radiocarbon | pable of testing the overkill hypothesis, but of posing legitimate temporal frequency distributions challenges to other extinction hypotheses, with the possible ex- ception of multifactor models that also invoke “first contact” ust over 42 y ago, Paul Martin (1) proposed that humans effects, such as the keystone herbivore (11), habitat modification ANTHROPOLOGY Jentered the contiguous United States via the Ice Free Corri- (12), and hyperdisease hypotheses (13). dor at ∼13,500 BP and there encountered almost three dozen We use two analytical techniques to identify dates of initial genera of now-extinct megafaunal mammals. Hunting of these megafaunal decline: for technical precision, an approach based on naïve prey fueled rapid human population growth, he argued, spacings (time lags between consecutive ordered dates), and, for resulting in both the colonization of the landmass stretching from simple understanding, an approach based on histograms of ob- the southern terminus of the North American ice sheets to the far served dates. The key property of spacings is that mean spacings tip of South America in 1,000 y, and the extinction of the mam- moths, mastodons, camels, horses, ground sloths, and other large Significance mammal taxa that had inhabited the Western Hemisphere for hundreds of thousands to millions of years before human arrival. Coincident with the human colonization of the Western Hemi- Martin (p. 973) closed his paper with the statement, “Should the sphere, dozens of genera of Pleistocene megafauna were lost to model survive future findings, it will mean that the extinction extinction. Following Martin, we argue that declines in the record of radiocarbon dates of extinct genera may be used as an in- chronology of the Pleistocene megafauna can be used to map the Homo sapiens ” dependent means of detecting the first presence of humans in spread of through the New World. the New World. Our results, based on analyses of radiocarbon Central to this hypothesis is the idea that small numbers of dates from Eastern Beringia, the contiguous United States, and humans can have large ecological impacts and that those impacts South America, suggest north to south, time, and space trans- should be directly observable in the paleontological record. If we gressive declines in megafaunal populations as predicted by the accept that premise as true, then as Martin argues, it is possible to overkill hypothesis. This finding is difficult to reconcile with other assess the timing of human arrival independently of direct ar- extinction hypotheses. However, it remains to be determined chaeological evidence by examining when megafaunal decline oc- whether these findings will hold with larger samples of radio- curred across time and space. In this paper, we use databases of carbon dates from all regions. radiocarbon dates on extinct megafauna from Eastern Beringia (EB), the contiguous United States (CUSA), and South America Author contributions: T.A.S. and S.R.P. designed research; T.A.S. and S.R.P. performed research; T.A.S., S.R.P., R.A.-S., and A.D.M. analyzed data; R.A.-S. and A.D.M. contributed (SA) to estimate the timing of initial population declines that ul- new reagents/analytic tools; and T.A.S., S.R.P., R.A.-S., and A.D.M. wrote the paper. timately resulted in extinction. We intend this exercise to be both a The authors declare no conflict of interest. direct test of the timing of extinction as proposed by Martin (1) and This article is a PNAS Direct Submission. Y.M. is a guest editor invited by the Editorial as an independent means of estimating the timing of human arrival Board. to each region. Similar approaches using paleoecological proxy 1To whom correspondence should be addressed. Email: [email protected]. records as possible indicators of human arrival have been applied This article contains supporting information online at www.pnas.org/lookup/suppl/doi:10. elsewhere, particularly on islands (2–7). 1073/pnas.1504020112/-/DCSupplemental. www.pnas.org/cgi/doi/10.1073/pnas.1504020112 PNAS Early Edition | 1of6 Downloaded by guest on October 2, 2021 are inversely proportional to population levels. For spacings, in- One pattern that emerges from comparative analyses of these dividual spacing values are regressed onto the midtime (ti+1 + ti)/2 three regions is that the shape of the EB frequency distribution is of the spacing interval using a generalized linear model with es- distinct from those of the CUSA and SA (Fig. 2). Given the timated breakpoints (representing times of onset of extinction). progressive loss of materials through time, radiocarbon ages are Decline dates are identified as the time before extinction for which expected to decrease nonlinearly with age from recent modes, waiting times between dates begin to increase, implying population producing a heavily right-skewed distribution (15, 16). Even after declines. This method produces both a likely date for extinction correction for taphonomic bias, the CUSA and SA datasets form onset and an associated approximate 95% CI (confidence interval). curves typical of this phenomenon. Potentially, loss should be For histogram creation, we first apply a jackknife method to identify even greater for these datasets than for others due to these dates optimal binning parameters for histogram creation (14) to create having been derived from bone and organic materials (e.g., dung), frequency distributions of calibrated radiocarbon dates, which are which may progressively be lost to weathering and/or dissolution then corrected for taphonomic bias following Surovell et al. (15). even if site sediments are not lost to erosion. Quite distinctly, there Taphonomic correction adjusts frequency distributions of radiocar- appears to be considerably less loss of bone through time in the EB dataset. We expect that this is due to a combination of ex- bon dates to account for the loss of sedimentary contexts through “ ” time due to erosion and weathering. Decline dates are estimated as cellent bone preservation in periodically frozen muck deposits occurring within the last mode before extinction. Spacings are also (17) and the sampling of extensive exposures of Pleistocene de- taphonomically corrected (SI Materials and Methods). posits due to gold mining activities (18). This pattern suggests that Both spacings and frequency analyses rest on three assump- unique taphonomic corrections are preferable for each region, but natural cycles in populations and sampling concerns are con- tions: (i) our radiocarbon datasets are representative of the rela- founding factors that make such unique corrections impracticable. tive frequencies of megafauna in the paleontological records of However, it is important to note that the taphonomic corrections each region; (ii) after taphonomic correction, temporal frequency we applied have little effect on estimating dates of megafaunal distributions positively correlate with population densities of iii decline (Fig.
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