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A Quantitative Model for Distinguishing Between Climate Change, Human Impact, and Their Synergistic Interactions As Drivers of T See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/295919520 A quantitative model for distinguishing between climate change, human impact, and their synergistic interactions as drivers of the late Quaternary megafaunal extinctions Article · October 2015 READS 143 4 authors, including: Emily Lindsey Natalia A. Villavicencio University of California, Berkeley University of California, Berkeley 15 PUBLICATIONS 735 CITATIONS 5 PUBLICATIONS 13 CITATIONS SEE PROFILE SEE PROFILE Anthony D Barnosky University of California, Berkeley 64 PUBLICATIONS 3,182 CITATIONS SEE PROFILE Available from: Emily Lindsey Retrieved on: 02 August 2016 A QUANTITATIVE MODEL FOR DISTINGUISHING BETWEEN CLIMATE CHANGE, HUMAN IMPACT, AND THEIR SYNERGISTIC INTERACTION AS DRIVERS OF THE LATE QUATERNARY MEGAFAUNAL EXTINCTIONS CHARLES R. MARSHALL,1 EMILY L. LINDSEY,1 NATALIA A. VILLAVICENCIO,1 AND ANTHONY D. BARNOSKY2 1Department of Integrative Biology and Museum of Paleontology, University of California, Berkeley, Berkeley, CA 94720 USA <[email protected]> 2Department of Integrative Biology, Museum of Paleontology, and Museum of Vertebrate Zoology, University of California, Berkeley, Berkeley, CA 94720 USA ABSTRACT.—A simple quantitative approach is presented for determining the relative importance of climate change and human impact in driving late Quaternary megafaunal extinctions. This method is designed to determine whether climate change or human impact alone can account for these extinctions, or whether both were important, acting independently (additively) and/or synergistically (multiplicatively). This approach is applied to the megafaunal extinction in the Última Esperanza region of southern Chile. In this region, there is a complex pattern of extinction. Records of environmental change include temperature proxies and pollen records that capture the transition from cold grasslands to warmer, moister forests, as well as evidence of initial human arrival. Uncertainty in extinction times and time of human arrival complicates the analysis, as does uncertainty about the size of local human populations, and the nature, strength, and persistence of their impacts through the late Pleistocene and early Holocene. Results of the Última Esperanza analysis were equivocal, with evidence for climate- and human-driven extinction, with each operating alone or additively. The results depend on the exact timing of extinctions and human arrival, and assumptions about the kinds of pressures humans put on the megafauna. There was little evidence for positive synergistic effects, while the unexpected possibility of negative synergistic interactions arose in some scenarios. Application of this quantitative approach highlights the need for higher precision dating of the extinctions and human arrival, and provides a platform for sharpening our understanding of these megafaunal extinctions. INTRODUCTION Alroy, 2001; Koch and Barnosky, 2006; Turvey, 2009), while in other areas, it appears that climate The cause(s) of the late Quaternary megafaunal change likely triggered extinction of some taxa extinctions continue to generate debate about the (Barnosky, 1985, 1986; Stuart et al., 2002, 2004; relative roles of humans (Martin, 1966, 1967, Guthrie, 2003, 2006). The case has also been 1973, 1984, 1990; Alroy, 1999, 2001; Flannery made for a “one-two punch” of human impact and and Roberts, 1999; Roberts et al., 2001), climate climate change exacerbating extinction intensity (Graham and Lundelius, 1984; Grayson, 1984, (Barnosky et al., 2004; Koch and Barnosky, 2006; 2001; Stuart, 1999; Stuart et al., 2002, 2004; Barnosky and Lindsey, 2010; Brook and Grayson and Meltzer, 2003; Trueman et al., 2005; Barnosky, 2012; Villavicencio et al., 2015). Wroe et al., 2006, 2013), and possible synergy However, few studies have attempted to between the two (Guilday, 1967, 1984; Barnosky quantitatively assess the relative importance of et al., 2004; Guthrie, 2006; Koch and Barnosky, humans, climate change, and/or interactions 2006; Barnosky and Lindsey, 2010; Brook and between the two in determining extinction Barnosky, 2012) in triggering extinctions. There is intensity. Moreover, when interaction effects are strong evidence that humans played a key role in postulated, there is seldom an effort to distinguish megafaunal extinctions in many geographic areas between additive effects—that is, climate change (MacPhee, 1999; Martin and Steadman, 1999; accounting for the extinction of some taxa and In: Earth-Life Transitions: Paleobiology in the Context of Earth System Evolution. The Paleontological Society Papers, Volume 21, P. David Polly, Jason J. Head, and David L. Fox (eds.). The Paleontological Society Short Course, October 31, 2015. Copyright © 2015 The Paleontological Society. MARSHALL ET AL.: QUANTIFYING CAUSES OF MASS EXTINCTION humans accounting for others—versus true flourishes under warmer conditions)? How did synergy, where interaction between human impact vegetation change affect species abundances, and climate change multiplies the extinction species interactions, and the ability of humans to intensity beyond the simple additive effects of the interact with them? To what extent did two factors acting independently. megafaunal change feed back onto vegetation Here, we present a new quantitative approach change, via the process of defaunation (Gill et al., designed to determine additive versus synergistic 2009, 2012; Galetti and Dirzo, 2013; Young et al., roles of human impact and climate change in 2013, 2014; Dirzo et al., 2014; Gill, 2014; driving extinction. The method ascribes Barnosky et al., 2015)? How did climate change, probabilities to the independent roles of human human activity, and megafaunal extinctions affect impact, non-human environmental change, and local fire regimes, and vice versa (Bond, 2005; interactions between the two in explaining the Bond and Keeley, 2005)? How did human- and observed temporal pattern of local extinction in a climate-driven changes elsewhere in South specified geographic region. The method is America impact the Última Esperanza region? described and applied to the Última Esperanza Beyond these fundamental considerations, region (Villavicencio et al., 2015), located in the shortcomings in the fossil record need to be taken Patagonia region of southern Chile. into account. For example, given the incompleteness of the record, when did the WHAT SIMPLE QUANTITATIVE MODELS various taxa actually become locally extinct? CANNOT CAPTURE When did humans first arrive? Moreover, to what extent might changes in the behaviors of taxa give While the fossil and paleoenvironmental records the false appearance of extinction? For example, of Última Esperanza are relatively rich (see in the study area, does the early disappearance of below), developing a comprehensive model of the fossil big cats indicate their extinction from the factors responsible for the extinction of the area, or simply that they moved elsewhere when megafauna in this, or any other region, is almost humans began to utilize caves that big cats had impossible. Such a model would require detailed formerly found hospitable, and that were a understanding of how the number of humans favorable depositional environment for present changed over time, whether they were fossilization? permanent residents in the region, what their hunting preferences and practices were, how the A SIMPLE QUANTITATIVE METHOD FOR various taxa responded to their presence (e.g., ASSESSING CAUSAL AGENTS OF whether after first contact some species learned to EXTINCTION AND THEIR POTENTIAL be wary of humans), what the interactions INTERACTION between the various taxa were and how these might have changed after human contact, among Given the lack of information needed to develop a many other questions. full mechanistic model of the megafaunal Quantifying non-human effects is equally extinctions, we present a simple correlative difficult. For example, in the focal region of approach that can distinguish between human southwestern Patagonia, while climate warming is impact, environmental change, and any thought to be the primary driver of environmental interactions between the two, in accounting for change as the region came out of the last ice age, the late Quaternary extinctions. This new method how did changes in climate actually translate into is designed to make use of available fossil and biotic transformations that might lead to environmental data, and accommodates the extinction? To what extent was the observed incompleteness of the fossil record. It requires temperature increase significant? How did rising input of three basic kinds of data: 1) the extinction sea level from the melting ice impact the region times of the megafauna, derived ideally from (particularly in the study area of southern multiple radiocarbon dates on multiple specimens; Patagonia)? Did rainfall change in any significant 2) a quantitative time series that records way, either seasonally or annually? Are indirect environmental change, such as pollen percentages effects of climate change more important than of environmentally sensitive plants, or direct changes in temperature or precipitation temperature proxies, such as those that come from (e.g., by triggering the replacement of cold- stable isotopes; and 3) some measure of probable adapted grasslands by Nothofagus forest, which human impact; for example, numbers of 2 THE PALEONTOLOGICAL SOCIETY PAPERS, V. 21 archaeological sites distributed
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