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Using Ecological Niche Modeling for Quantitative Biogeographic Analysis: a Case Study of Miocene and Pliocene Equinae in the Great Plains

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Using Ecological Niche Modeling for Quantitative Biogeographic Analysis: a Case Study of Miocene and Pliocene Equinae in the Great Plains Paleobiology, 35(4), 2009, pp. 587–611 Using ecological niche modeling for quantitative biogeographic analysis: a case study of Miocene and Pliocene Equinae in the Great Plains Kaitlin Clare Maguire and Alycia L. Stigall Abstract.—The subfamily Equinae in the Great Plains region of North America underwent a dramatic radiation and subsequent decline as climate changed from warm and humid in the middle Miocene to cooler and more arid conditions during the late Miocene. Here we use ecological niche modeling (ENM), specifically the GARP (Genetic Algorithm using Rule-set Prediction) modeling system, to reconstruct the geographic distribution of individual species during two time slices from the middle Miocene through early Pliocene. This method combines known species occurrence points with environmental parameters inferred from sedimentological variables to model each species’ fundamental niche. The geographic range of each species is then predicted to occupy the geographic area within the study region wherever the set of environmental parameters that constrain the fundamental niche occurs. We analyze changes in the predicted distributions of individual species between time slices in relation to Miocene/Pliocene climate change. Specifically, we examine and compare distribution patterns for two time slices that span the period from the mid-Miocene (Barstovian) Climatic Optimum into the early Pliocene (Blancan) to determine whether habitat fragmentation led to speciation within the clade and whether species survival was related to geographic range size. Patchy geographic distributions were more common in the middle Miocene when speciation rates were high. During the late Miocene, when speciation rates were lower, continuous geographic ranges were more common. Equid species tracked their preferred habitat within the Great Plains region as well as regionally throughout North America. Species with larger predicted ranges preferentially survived the initial cooling event better than species with small geographic ranges. As climate continued to deteriorate in the late Miocene, however, range size became irrelevant to survival, and extinction rates increased for species of all range sizes. This is the first use of ENM and GARP in the continental fossil record. This powerful quantitative biogeographic method offers great promise for studies of other taxa and geologic intervals. Kaitlin Clare Maguire. Department of Integrative Biology, University of California, Berkeley, California 94720. E-mail: [email protected] Alycia L. Stigall. Department of Geological Sciences and OHIO Center for Ecology and Evolutionary Studies, Ohio University, Athens, Ohio 45701. E-mail: [email protected] Accepted: 9 February 2009 Introduction mosaic of savanna and riparian forests that The subfamily Equinae underwent a major supported browsing, grazing, and mixed- radiation during the Miocene from one feeding horses (Webb 1983). Following the species, Parahippus leonensis, to 70 species, radiation, members of the Equinae exhibit a reaching its highest diversity during the variety of dental and muzzle morphologies as middle Miocene (Clarendonian) with 13 well as a diverse range of body sizes and limb named genera (MacFadden 1992; Hulbert proportions (MacFadden 1992). It has been 1993) (Fig. 1). The main interval of equid previously hypothesized that this morpho- radiation coincides with the first major logical diversity arose as equids with over- cooling event of the Miocene (Zachos et al. lapping distributions partitioned their niches 2001), and the equid radiation has been to exploit different resources as climate classically attributed to an adaptive response changed (MacFadden 1992; Webb et al. following climate and vegetation change 1995). This diversity peak, however, did not during the Miocene (MacFadden and Hulbert last. The equid clade began to decline during 1988; Hulbert 1993). As the climate became the late Miocene, and by the middle of the cooler and drier during the Miocene, vegeta- Pliocene only three genera were extant (Hul- tion shifted from woodland dominated to a bert 1988). Competition with other grazers ’ 2009 The Paleontological Society. All rights reserved. 0094-8373/09/3504–0007/$1.00 588 KAITLIN C. MAGUIRE AND ALYCIA L. STIGALL FIGURE 1. Temporally calibrated cladogram of Equinae modified from Maguire and Stigall (2008). Time slice divisions are indicated at the top. Narrow horizontal lines indicate ghost lineages, whereas bold horizontal lines indicate the recorded range. Thick gray lines indicate species whose ranges were modeled in this analysis. ECOLOGICAL NICHE MODELING OF EQUINAE 589 (e.g., artiodactyls, rodents) was previously Although the diversification of the Equinae inferred to have caused the decline of the is hypothesized to have resulted from climate clade (Simpson 1953; Webb 1969, 1984; Stan- change and ecological specialization, the ley 1974), but it is now attributed to the interaction between individual species and continuation of cooling temperatures and environmental conditions has not previously more arid climatic conditions (MacFadden been quantitatively analyzed. Here we recon- 1992; Webb et al. 1995). struct the geographic ranges for individual Most paleobiogeographic studies of equids species of Miocene to early Pliocene horses in have focused on large-scale patterns, such as concert with environmental conditions to migration patterns from North America to the assess how changing climate and shifting Old World (e.g., MacFadden 1992). Other habitats affected evolutionary patterns in the paleobiogeographic studies have grouped Equinae. In order to study the distribution of equids with other Miocene mammalian clades horses relative to ecological changes, we use to demonstrate general patterns, such as ecological niche modeling (ENM). A species’ retreat to the Gulf Coast region as the climate ecological, or fundamental, niche is defined as became cooler and more arid in northern areas the set of environmental tolerances and limits of North America during the late Miocene in multidimensional space that defines where (e.g., Webb 1987; Webb et al. 1995). A more a species is potentially able to maintain focused study examining the distribution of populations (Grinnell 1917; Hutchinson individual equid species can provide a frame- 1957). Determining the ecological niche of a work in which to analyze habitat fragmenta- taxon is essential in determining its potential tion, niche partitioning, and speciation. Be- distribution (Peterson 2001) as well as taxon/ cause the radiation is hypothesized to have environment interactions. A species may resulted from climate change, an integrated potentially occupy the entire geographic assessment of the ecology and biogeography is region in which its ecological niche occurs; necessary to fully understand evolutionary although biotic interactions, including com- patterns in the clade. Shotwell (1961) recog- petition and predation may limit a species nized this relationship in his study on the range to a smaller ‘‘realized niche’’ (Lomolino biogeography of horses in the northern Great et al. 2006). Because determination of these Basin, in which he observed changes in species biotic interactions from the fossil record is composition that coincided with vegetation rarely possible, we turn here instead to change. Shotwell concluded that this shift modeling species’ ecological niches. resulted from immigration of equid species Ecological niche modeling is a GIS-based adapted to the new vegetative regime from method in which species’ environmental other regions into the Great Basin. tolerances are modeled on the basis of a set Recent advances in understanding the of environmental conditions at locations phylogenetic relationships provide a more where the species is known to occur. The robust framework for new analyses. Further- species range is then predicted to occupy more, numerous recent studies have investi- areas where the same set of environmental gated the environmental and climatic condi- conditions occurs in the study region. Funda- tions of the Great Plains during the Miocene. mentally, this approach directly utilizes sed- These studies comprise analyses of vegetation imentary data to determine the likely distri- type (e.g., Fox and Koch 2003; Stro¨mberg bution of taxa, thus providing a framework to 2004; Thomasson 2005), paleosol composition examine interactions between abiotic and (e.g., Retallack 1997, 2007), and climate biotic factors that control species distribution proxies including stable oxygen and carbon and how these shift through time. ENM- isotopes of foraminifera from deep-sea cores based range prediction provides a framework (e.g., Woodruff et al. 1981; Zachos et al. 2001) to test hypotheses about similarities/differ- that provide a rich source of environmental ences in evolutionary and/or biogeographic information previously unavailable for equid patterns exhibited as climate and environ- ecological biogeographic analyses. ments shifts through time (Stigall and Lieber- 590 KAITLIN C. MAGUIRE AND ALYCIA L. STIGALL man 2006; Stigall 2008). ENM methods were constrain the rise and fall of the Equinae in originally developed for modern taxa, and relation to environmental and climatic high levels of predictive accuracy have been change. We model ecological niches and documented (Peterson and Cohoon 1999; predict geographic ranges in order to study Peterson 2001; Stockwell and Peterson 2002). how species’ distributions shifted through In particular, analyzing niche
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