(Microsyopidae, Primates) from the Paleocene-Eocene Thermal Maximum: One Species Or Two?
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Molar Size and Shape Variation in a Large Sample of Niptomomys (Microsyopidae, Primates) from the Paleocene-Eocene Thermal Maximum: One Species or Two? by Rosa S. Felibert Department of Anthropology University of Florida 2017 ABSTRACT The oldest euprimates first appear during a period of rapid, short-term, global warming ~56 mya known as the Paleocene-Eocene Thermal Maximum (PETM). Plesiadapiform primates of similar size and dental morphology to euprimates were present in North America before the PETM, and may have been affected by the arrival of euprimates as ecological competitors. Screenwashing PETM fossil localities in the Bighorn Basin, Wyoming, has yielded many fossils (N≈600) of the microsyopid plesiadapiform Niptomomys. N. doreenae is known from before and after the PETM and may range through it. A second taxon, N. favorum, characterized by its small size and squarer M2 occlusal outline, was described from the large Castle Gardens locality sample. To better characterize PETM primate diversity, we test the validity of N. favorum against a sample of Niptomomys from Castle Gardens, other PETM localities, and published measurements. M2 occlusal outlines (N=62) revealed a continuous range from square to lingually compressed that encompasses the holotype of N. favorum. Linear measurements of M1 (N=127) and M2 (N=163) indicated M1’s from Castle Gardens are larger than those of later PETM localities (p=0.038), but produced no outliers, suggesting the PETM fauna contains one species of Niptomomys. PETM lower molars are smaller than all other measured Niptomomys teeth, paralleling the response to warming effects recorded in larger-bodied mammal lineages. These results are consistent with either a single lineage of Niptomomys that became smaller during the PETM, or a small immigrant taxon (N. favorum) that was transitionally present in the Bighorn Basin during the PETM. 1 INTRODUCTION The Paleocene-Eocene Thermal Maximum (PETM) was a rapid warming event that occurred about 56 million years ago (mya), during which global temperatures increased ~5-8 degrees C in the span of 20,000 years or less. The entire event lasted 200,000 years and was caused by an increase in the amount of carbon dioxide (CO2) in the atmosphere (McInerney & Wing, 2011). This warming event facilitated the dispersal and dwarfing of mammals in North America, as well as the migration of euprimates, or primates of modern aspect (Bloch, Silcox, Boyer, & Sargis, 2007). The arrival of euprimates to North America may have caused ecological competition with endemic plesiadapiforms, a sister group of euprimates (Bloch, Silcox, Boyer, & Sargis, 2007; Gingerich, 2003). Adaptive changes may have occurred during the PETM to fill new and rapidly shifting ecological niches, and these provide the setting of this paper. One such adaptive change is body size. Many studies (Fleagle, 1978; Hutchinson & MacArthur, 1959; Newell, 1949; Cope, 1887; Gould, 1966; Gingerich & Smith, 1985; Peters, 1986; Maurer, Brown, & Rusler, 1992) have shown that body size is important when studying the ecology of past organisms, as well as when attempting to reconstruct an extinct taxon’s biology and life history. One such group of extinct organisms is the suborder of primitive primate-like mammals, Plesiadapiformes. Plesiadapiforms appear in the fossil record ~65 mya and have been argued to have shared a recent common ancestor with euprimates (Bloch, Silcox, Boyer, & Sargis, 2007). This group of mammals, which flourished during the Paleocene and Early Eocene of North America, shares numerous traits with living primates, even including a species with a nail instead of the more primitive claw (Bloch 2 & Boyer, 2002). Understanding plesiadapiforms is a crucial component to understanding primate evolution, and therefore, our own evolutionary history. The suborder Plesiadapiformes contains 12 families (Bloch, Silcox, Boyer, & Sargis, 2007; Silcox, 2001; Silcox & Gunnell, 2008). One of these is Microsyopidae, a diverse family of over 20 species that ranged from the Late Paleocene through Middle Eocene of North America (Fleagle, 2013; Gunnell, 1989; Silcox & Gunnell, 2008). Fossils of the microsyopid plesiadapiform genus Niptomomys have been found in North America ranging from before, during, and after the PETM. To date, there have been three species of Niptomomys identified: N. doreenae, N. thelmae, and the more elusive N. favorum. N. favorum from the Castle Gardens locality in the Bighorn Basin, Wyoming, was diagnosed based on an isolated M1/2 (UCMP 212635) as being smaller and squarer in occlusal outline than N. doreenae and N. thelmae, (Strait, 2001). The extant primate literature has shown that there is a high correlation between tooth size, specifically lower cheek teeth, and body size (Gingerich, 1974; Gingerich & Schoeninger, 1979; Gingerich & Smith, 1985; Gingerich, Smith, & Rosenberg, 1982; Pilbeam & Gould, 1974). This study assessed the occlusal outline of upper molars from Castle Gardens and measured a large sample of M1 and M2 Niptomomys specimens to determine whether N. favorum can be identified as a separate species existing alongside N. doreenae during the PETM, or if specimens attributed to N. favorum from Castle Gardens are merely a manifestation of dwarfing with a continuous N. doreenae lineage through the PETM, as has been observed in other PETM lineages (Gingerich, 2003; Secord et al., 2012). 3 LITERATURE REVIEW The Paleocene-Eocene Thermal Maximum The PETM was a massive, rapid global warming event that caused temperatures to rise ~5-8 degrees Celsius at a rapid rate and persist for a span of ~200,000 years. There have been numerous contributions made to the literature about the PETM, its onset, and its ecological consequences for mammals. McInerney and Wing (2011) thoroughly discussed changes in the carbon cycle, precipitation, ecosystems, and temperature during the PETM, and compared its effects to the effects that similar, even more rapid, anthropogenic global warming event could have in the future. Chester and colleagues (2010) determined that rapid morphological change in carnivorous species occurred during the PETM as a way of adapting to the changing climate and the availability of food. This conclusion was based on Rosenzweig’s (1968) study on Actual Evapotranspiration (AE), the amount of water that is evaporated from soil and transpired from plants into the atmosphere, and its relationship with carnivore body size. AE is correlated with the availability of water and solar energy and is therefore a good predictor of mammalian carnivore productivity levels. Rosenzweig found that environments lacking either or both of these resources are scarce in food, which he found limited the body size of mammalian carnivores (Rosenzweig, 1968). Chester and colleagues (2010) interpreted this research and concluded that selective pressures, such as lack of water or increased temperature, could cause smaller body sizes in mammals during the PETM. Gingerich (2003) also related mammal body size to the PETM. His research showed that mammalian body size dwarves in the same strata where the PETM occurred, providing evidence that climate change and faunal size change are related. 4 He posited that two different types of changes in body size occurred: transient and permanent. A transient change in body size is especially interesting to study because evidence shows body size decreasing during the PETM and returning to pre-PETM size after the PETM. Gingerich (2003) posited that these adaptive changes had to happen for immigrant taxa, which could only migrate due to the conditions the PETM caused, to coexist with endemic taxa. Secord and colleagues (2012) related the PETM to mammal body size by discussing the change of body size in equids during and after the PETM. This was determined by examining the isotopic composition of mammal teeth and finding that body size in equids was negatively correlated with the temperatures inferred from the oxygen isotopes in those teeth. At the carbon isotope excursion (CIE) onset, equids had a 30% size decrease compared to a 76% size increase at the end of the CIE. This decrease in body size may have been caused by higher temperatures and increased CO2 concentrations in the atmosphere. This paper also depicts the correlation between the decrease of carbon-13 and the increase of oxygen-18 during the PETM, and suggests that environmental and faunal changes during the PETM occurred extremely rapidly. Importance of Body Size in Ecology Several studies have demonstrated the importance of body size in terms of ecology and macro- and microevolution (Fleagle, 1978; Hutchinson & MacArthur, 1959; Newell, 1949; Cope, 1887; Gould 1966; Speakman, 2005; Peters & Wassenberg, 1983; Schmidt-Nielsen, 1984; Kelt & Van Vuren, 1999; Savage et al., 2007; Duncan, Forsyth, & Hone, 2007; Capellini, Venditti, & Barton, 2010) 5 Peters (1983) provided a comprehensive book explaining the ways in which organism body size affects ecology. On a fundamental level, body size affects a number of physiological functions, such as metabolism, heat production and heat loss, rate of energy use, and dynamics of locomotion. He discussed that these effects on physiological function are the reason for the wide range of animal diversity on the planet. Schmidt-Nielsen (1984) also provided a comprehensive introduction to the topic of scaling and its importance in ecology. He described the concept of food extraction, the amount of food resources an animal consumes and how this directly affects population densities