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Understanding Eocene Primate Palaeobiology Using a Comprehensive Analysis of Living Primate Ecology, Biology and Behaviour

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Understanding Eocene Primate Palaeobiology Using a Comprehensive Analysis of Living Primate Ecology, Biology and Behaviour Palaeobio Palaeoenv (2012) 92:573–583 DOI 10.1007/s12549-012-0089-1 ORIGINAL PAPER Understanding Eocene primate palaeobiology using a comprehensive analysis of living primate ecology, biology and behaviour Michelle L. Sauther & Frank P. Cuozzo Received: 31 March 2012 /Revised: 28 May 2012 /Accepted: 6 June 2012 /Published online: 7 July 2012 # Senckenberg Gesellschaft für Naturforschung and Springer 2012 Abstract The comparative method is central to interpreta- assumption that heavy tooth wear leads to the rapid death tions of Eocene primate palaeobiology. This method rests of the individual. Finally, our data on trauma and injury upon a thorough study of analogous living forms. With a from a living lemur population suggest that the reported rapidly increasing knowledge of such forms, most notably wrist injury in Darwinius masillae (i.e. “Ida”)didnot the Malagasy lemurs, our ability to advance the study of necessarily lead to her death, as numerous ring-tailed Eocene primate ecology, biology and behaviour far lemurs at Beza Mahafaly survive with similar or even exceeds that of even just a few years ago. Here we present more traumatic injuries and maintain the ability to climb. such a comparison. Based on our data collected from both Thus, our data from living primates provide a broad com- living lemurs and extant lemur skeletal specimens, we are parative framework for interpreting the ecology, biology able to make a number of comparisons that provide insight and behaviour of Eocene forms. into middle Eocene primate ecology and palaeobiology. At the Beza Mahafaly Special Reserve, Madagascar, omnivo- Keywords Lemur . Dental ecology . Notharctus . Darwinius . rous living ring-tailed lemurs that feed on large, hard and Laterality tough fruits display a pattern of frequent post-canine tooth wear laterality (62 %) when compared to sympatric, folivorous Verreaux’s sifaka (4 %). Our results indicate that Introduction Notharctus does not display a high frequency of tooth wear laterality (7 %), indicating folivory without processing large, There has been a long history of interpreting the ecology hard fruits with its postcanines. Our data on Notharctus tooth and behaviour of extinct primates through comparisons with wear also indicate, similar to living ring-tailed lemurs at Beza comparable, living forms (Covert 1986, 1997), and the Mahafaly, that numerous individuals (21 %) survived long comparative method is central for answering many ques- enough to experience heavy tooth wear, contrary to the tions in primate evolutionary biology (Anthony and Kay 1993). Specifically, for Eocene primates, the extant strepsir- rhine primates have provided an important template for This article is a contribution to the special issue “Messel and the terrestrial Eocene—Proceedings of the 22nd Senckenberg Conference” interpreting body size, anatomy, locomotion and dietary preferences for these extinct forms (Covert 1986). These * : M. L. Sauther ( ) F. P. Cuozzo early comparisons focused primarily on large questions of Department of Anthropology, University of Colorado, Campus Box 233, Boulder, CO 80309-0233, USA overall primate adaptations (e.g. Covert 1986)andwere e-mail: [email protected] limited in terms of interpreting their ecology. This approach F. P. Cuozzo was primarily due to the paucity of studies of extant strep- e-mail: [email protected] sirrhine behavioural ecology that could provide good living models to assess comparable extinct forms. However, over F. P. Cuozzo the past two decades, there has been a dramatic increase in Department of Anthropology, University of North Dakota, 236 Centennial Drive, Stope 8374, research on extant lemur ecology, biology and locomotion Grand Forks, ND 58202, USA (e.g. Gould and Sauther 2006; Jolly et al. 2006; Kappeler 574 Palaeobio Palaeoenv (2012) 92:573–583 and Ganzhorn 1993; Rakotosamimanana et al. 1999). With Methods continued field work by workers from various institutions (e.g. University of Michigan, University of Colorado, Uni- Extant samples versity of California–Berkeley, the Carnegie Museum of Natural History, the Denver Museum of Nature and Sci- Within the framework of our ongoing Lemur Biology Project, ence), which has produced large samples of Eocene primate we have collected in-depth dental and biological data for a materials from the intermontane basins of North America wild population of lemurs living within and around the Beza (e.g. Anemone and Covert 2000; Beard et al. 1991; Beard Mahafaly Special Reserve (BMSR), in southwestern Mada- and MacPhee 1994; Covert and Hamrick 1993; Gunnell gascar. As part of a project to understand and monitor the 1995, 1997; Murphey et al. 2001; Rose et al. 1999; Ross effects of habitat and climate on lemur ecology, over 300 and Covert 2000; Williams and Covert 1994), it is now individual ring-tailed lemurs, Lemur catta, have been moni- possible to provide important contextual data for expanding tored over a 25-year period at BMSR. The collected data our understanding of the biology of extinct Eocene (and include health and medical evaluations as well as dental other) fossil primates. assessments of these individuals in yearly captures (Cuozzo Assessing patterns of tooth-use wear in primate fossils et al. 2010; Sussman et al. 2012) across a 10-year period has a long history. However, since its development in the beginning in 2003. In 2006 we also collected comparable 1970s and 1980s (e.g. Walker et al. 1978), analyses of biological and dental information from 25 living ring-tailed dental microwear have dominated dental wear assessment, lemurs at Tsimanampesotse National Park, in southwestern with patterns of overall or gross wear receiving far less Madagascar. In addition, skeletal remains for wild ring-tailed attention (e.g. Ungar 2002). Our long-term work in Mada- lemurs (L. catta) and sifaka (Propithecus verreauxi)havebeen gascar combines data on feeding ecology and food prop- collected over a 25-year period, now housed at the BMSR, erties in a single primate community to identify the and these provide patterns of dental ecology and pathology as proximate causes of primate tooth wear, and thus provides well as skeletal injury, disease and trauma. Extant comparative an interpretive framework for assessing patterns of gross lemur dental and skeletal samples also come from the Amer- tooth wear in primate fossils. Specifically, we have docu- ican Museum of Natural History (AMNH), the United States mented identifiable patterns of tooth wear produced by National Museum of Natural History (USNM), The Natural consuming specific foods with particular properties (e.g. History Museum, London, Harvard’sMuseumofCompara- Cuozzo and Sauther 2004, 2006; Sauther and Cuozzo tive Zoology and the Museum für Naturkunde, Berlin 2009). In numerous descriptions of Eocene primates (BMNH). Quantitative data on extant ring-tailed lemur tooth (and many other early Tertiary mammals), specimens are wear from BMSR presented herein come from individuals often noted as displaying varying degrees of gross tooth captured in 2011 (n051). The 2011 data set includes lemurs wear, with authors commonly lamenting that “heavy” ranging in age from 3 years through those in their mid teens. wear obscures dental traits needed for taxonomic identifi- These older lemurs were first captured in 2003 as full adults, cation. Yet these patterns of heavy tooth wear can be quite and thus they were categorised by their minimum age when informative in terms of assessing the behaviour and ecol- captured in 2011. All animal captures were conducted with ogy of extinct forms (e.g. Jablonski et al. 2008). IACUC (Institutional Animal Care and Use Committee) ap- In this paper we use data collected during our 25-year proval from the University of North Dakota and/or the Uni- research on living lemur’s dental ecology and biology and versity of Colorado–Boulder and followed protocols from our long-term work on extant lemur skeletal samples established over the 25-year history of this project (e.g. to contextualise Eocene primate ecology, biology and be- Sauther and Cuozzo 2009; Sussman et al. 2012). haviour (Cuozzo and Sauther 2004, 2006; Sauther et al. 2002; Sauther and Cuozzo 2008). Specifically, we compare Eocene notharctine samples our extant dental and biological data to information de- rived from our analyses of 29 notharctine specimens col- Notharctines have long been argued to share a number of lected from a number of museums in North America. We morphological traits with living Malagasy indriids, especial- use these data to explore patterns of tooth wear as well as ly members of the genus Propithecus (Covert 1986). Based the potential of laterality in tooth wear (or lack thereof) in on their anatomy, notharctines are believed to have been living and/or extant lemurs, in which tooth wear laterality folivorous leapers similar to living indriids such as sifaka. corresponds to broad dietary behaviours and categories. For example, the intermembral index of Notharctus is 60, We also use our extant primate biological information to which is similar to the intermembral index of 59 found in the re-visit the interpretation of the taphonomy and behaviour vertical clinger and leaper P. verreauxi (Fleagle 1998). Here of one of the most important Eocene primate finds to date, we present tooth wear and wear laterality data for 29 Ida (Darwinius masillae). Notharctus specimens (Table 1), obtained from collections Palaeobio Palaeoenv (2012) 92:573–583 575 housed at the USNM, the Denver Museum of Nature and number of tooth positions scored. A first molar mean wear Science (DMNS) and the AMNH. score was also calculated, as follows: combined M1 and m1 wear scores/total number of first molar tooth positions Wear scores scored for each individual. These indices were divided into the following wear score categories: “low”0scores 0.00– Each tooth position was scored for degree of overall or gross 1.00; “medium”0scores 1.01–2.00; “medium high”0scores wear in both the Notharctus and L. catta samples. Wear 2.01–3.00 and “high”03.01–4.00.
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