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Evolution of the Earliest Horses Driven by Climate Change in the Paleocene-Eocene Thermal Maximum

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Evolution of the Earliest Horses Driven by Climate Change in the Paleocene-Eocene Thermal Maximum University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln Earth and Atmospheric Sciences, Department Papers in the Earth and Atmospheric Sciences of 2-24-2012 Evolution of the Earliest Horses Driven by Climate Change in the Paleocene-Eocene Thermal Maximum Ross Secord University of Nebraska-Lincoln, [email protected] Jonathan I. Bloch Florida Museum of Natural History, University of Florida, [email protected] Stephen G. B. Chester Yale University, [email protected] Doug M. Boyer Brooklyn College, City University of New York, [email protected] Aaron R. Wood South Dakota School of Mines and Technology, [email protected] See next page for additional authors Follow this and additional works at: https://digitalcommons.unl.edu/geosciencefacpub Part of the Earth Sciences Commons Secord, Ross; Bloch, Jonathan I.; Chester, Stephen G. B.; Boyer, Doug M.; Wood, Aaron R.; Wing, Scott L.; Kraus, Mary J.; McInerney, Francesca A.; and John Krigbaum, "Evolution of the Earliest Horses Driven by Climate Change in the Paleocene-Eocene Thermal Maximum" (2012). Papers in the Earth and Atmospheric Sciences. 310. https://digitalcommons.unl.edu/geosciencefacpub/310 This Article is brought to you for free and open access by the Earth and Atmospheric Sciences, Department of at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Papers in the Earth and Atmospheric Sciences by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. Authors Ross Secord, Jonathan I. Bloch, Stephen G. B. Chester, Doug M. Boyer, Aaron R. Wood, Scott L. Wing, Mary J. Kraus, Francesca A. McInerney, and John Krigbaum This article is available at DigitalCommons@University of Nebraska - Lincoln: https://digitalcommons.unl.edu/ geosciencefacpub/310 Published in Science vol. 335 no. 6071 (February 24, 2012), pp. 959-962; doi: 10.1126/science.1213859 Copyright © 2012 AAAS. Used by permission. Submitted September 12, 2011; accepted January 13, 2012. Evolution of the Earliest Horses Driven by Climate Change in the Paleocene-Eocene Thermal Maximum Ross Secord,1,2 Jonathan I. Bloch,2, Stephen G. B. Chester,3 Doug M. Boyer,4 Aaron R. Wood,5,2 Scott L. Wing,6 Mary J. Kraus,7 Francesca A. McInerney,8 John Krigbaum,9 1. Department of Earth and Atmospheric Sciences, University of Nebraska, Lincoln, NE 68588, USA 2. Florida Museum of Natural History, University of Florida, Gainesville, FL 32611–7800, USA 3. Department of Anthropology, Yale University, New Haven, CT, 06520, USA 4. Department of Anthropology and Archaeology, Brooklyn College, City University of New York, New York, NY 11210, USA 5. Department of Geology and Geological Engineering, South Dakota School of Mines and Technology, Rapid City, SD 57701, USA 6. Department of Paleobiology, Smithsonian Museum of Natural History, Washington, DC 20560, USA 7. Department of Geological Sciences, University of Colorado, Boulder, CO 80309, USA 8. Department of Earth and Planetary Sciences, Northwestern University, Evanston, IL 60208, USA 9. Department of Anthropology, University of Florida, Gainesville, FL 32611–7305, USA Corresponding author — R. Secord, email: [email protected] Abstract Body size plays a critical role in mammalian ecology and physiology. Previous research has shown that many mammals became smaller during the Paleocene-Eocene Thermal Maximum (PETM), but the timing and magni- tude of that change relative to climate change have been unclear. A high-resolution record of continental climate and equid body size change shows a directional size decrease of ~30% over the first ~130,000 years of the PETM, followed by a ~76% increase in the recovery phase of the PETM. These size changes are negatively correlated with temperature inferred from oxygen isotopes in mammal teeth and were probably driven by shifts in temper- ature and possibly high atmospheric CO2 concentrations. These findings could be important for understanding mammalian evolutionary responses to future global warming. nterest in how organisms respond at the onset of the CIE in 21 ky or less or the availability of food resources re- to climate change has intensified in (2), temperature increase was slower, lated to primary productivity (11). Berg- Irecent years with projected warm- peaking 60 ky or more into the CIE mann’s rule predicts that average mam- ing of ~2° to 4°C over the next century (4) at ~5° to 10°C above pre-CIE lev- malian body size should decrease with (1). Although models can be devel- els (5, 6). We use oxygen isotope values warming climate, and smaller size in en- oped to predict evolutionary responses in mammal teeth as a proxy for local dotherms has even been suggested as to warming of this magnitude, empiri- temperature change in the continen- a third “universal” response to warm- cal examples must be drawn from fos- tal interior of North America, and we ing, along with changes in phenology sil or historical records. Here we report show that equid body size during the and species distribution (10). Declining a dramatic example of shifts in body PETM was negatively correlated with body size has been attributed to warm- size in the earliest known horses (fam- temperature. ing over decadal and millennial scales ily Equidae) during the Paleocene-Eo- In extant mammals and birds (endo- in some living endotherms (12, 13), but cene Thermal Maximum (PETM) (~56 therms), closely related species or pop- many counterexamples also exist (10). million years ago). The PETM is rec- ulations within a species are gener- Furthermore, it is difficult to distinguish ognized in marine and continental re- ally smaller-bodied at lower latitudes, natural selection (genetic change) from cords by an abrupt negative carbon iso- where ambient temperature is greater ecophenotypic plasticity (morphologi- tope excursion (CIE) that lasted ~175 (7). This relationship, known as Berg- cal response not genetically fixed) over thousand years (ky), caused by the re- mann’s rule, is followed by ~65 to 75% such short time scales. The size change lease of thousands of gigatons of car- of studied extant mammals (8, 9). The documented here was, however, sus- bon to the ocean-atmosphere system (2, cause of Bergmann’s rule is usually at- tained over thousands of generations, 3). Some marine records suggest that tributed to thermoregulation and the strongly suggesting that natural selec- although δ13C values shifted rapidly optimization of body size (10) and/ tion was the cause. 959 960 S ECORD ET AL . IN S CIENCE 335 (2012) Previous studies lacked the strati- graphic resolution to recognize patterns in body size change within the PETM but demonstrated gross changes in size in several mammal lineages, based on first molar tooth area (14, 15). Size changes occurred in herbivorous ungulates (Pe- rissodactyla, Artiodactyla, Condylarthra, and Tillodontia), Primates, and fauni- vores and omnivores (Creodonta, Car- nivoramorpha, and Palaeanodonta), af- fecting both immigrant and endemic taxa (Figure 1). These changes conform well to Bergmann’s rule in terms of the ex- pected direction of size change. Quan- tifying published results, size reduc- tion occurred in 10 Paleocene genera that ranged into the PETM, represent- ing 38% of the range-through genera. This was followed by post-PETM size in- creases in eight of these genera, indicat- Figure 1. Summary of percent mean body size change in genera that exhibit change from ing that body size response was strongly the latest Paleocene to the PETM (left), and from the PETM to the post-PETM (right). No ge- taxon-specific (Figure 1 and table S7). nus exhibits a size increase in the PETM or a decrease after the PETM. Compiled from pub- Post-PETM size increases also occurred lished sources, except for Sifrhippus from this study. Asterisks indicate genera that first ap- pear in the PETM. See table S7 for a summary of all PETM taxa and sources. in an additional eight genera, seven of which first appeared in the PETM (Fig- ure 1). Together these 16 genera repre- enrichment (19). Plants in turn track the inversely proportional to the interval of 13 sent a size increase in 40% of PETM gen- δ C value of atmospheric CO2, with in- time over which rates are measured, be- era that ranged into post-PETM biozone fluences from environmental factors cause of the occurrence of small rever- Wa-1 (Figure 2). such as humidity and vegetation den- sals in the variable. The relationship be- Sifrhippus [formerly Hyracotherium sity (20, 21). At Cabin Fork, phenacodon- tween rates of change and the lengths of (16)] first appeared in North America tids (Ectocion and Copecion) record a neg- intervals over which they are observed 13 and Europe during the PETM. Because of ative shift of ~4.6 per mil (‰) in δ CE at is used to determine evolutionary mode the lack of a plausible ancestor on these the onset of the CIE (Figure 2A). This is (22). Our mwLRI results indicate with continents, it is widely thought to be an consistent with estimates of atmospheric 95% confidence that Sifrhippus body size immigrant that crossed high-latitude dis- change of ~4.6‰ during the PETM from directionally decreased from its first ap- persal routes opened by PETM warm- a leaf discrimination model (20) and pearance to the 41-m level, after which ing (17). We use Sifrhippus to document ~4.0‰ from modeling of marine carbon- stratigraphic resolution and sample sizes mammalian body size change within the ate dissolution (2), indicating that phe- are insufficient to distinguish between 13 PETM. Sifrhippus is the most abundantly nacodontid δ CE is primarily tracking directional and random evolutionary represented genus in new collections atmospheric δ13C values, rather than en- change. Thus, Sifrhippus experienced sus- from the Cabin Fork area (~10 km2) of vironmental change. tained selection for diminutive body size the southern Bighorn Basin, Wyoming, Sifrhippus sandrae first appears at for ~130 ky.
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