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Comparative Ecomorphology and Biogeography of Herpestidae and Viverridae (Carnivora) in Africa and Asia

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Comparative Ecomorphology and Biogeography of Herpestidae and Viverridae (Carnivora) in Africa and Asia Comparative Ecomorphology and Biogeography of Herpestidae and Viverridae (Carnivora) in Africa and Asia Gina D. Wesley-Hunt1, Reihaneh Dehghani2,3 and Lars Werdelin3 1Biology Department, Montgomery College, 51 Mannakee St. Rockville, Md. 20850, USA; 2Department of Zoology, Stockholm University, SE-106 91, Stockholm, Sweden; 3Department of Palaeozoology, Swedish Museum of Natural History, Box 50007, SE-104 05, Stockholm, Sweden INTRODUCTION Ecological morphology (ecomorphology) is a powerful tool for exploring diversity, ecology and evolution in concert (Wainwright, 1994, and references therein). Alpha taxonomy and diversity measures based on taxon counting are the most commonly used tools for understanding long-term evolutionary patterns and provide the foundation for all other biological studies above the organismal level. However, this provides insight into only a single dimension of a multidimensional system. As a complement, ecomorphology allows us to describe the diversification and evolution of organisms in terms of their morphology and ecological role. This is accomplished by using quantitative and semi-quantitative characterization of features of organisms that are important, for example, in niche partitioning or resource utilization. In this context, diversity is commonly referred to as disparity (Foote, 1993). The process of speciation, for example, can be better understood and hypotheses more rigorously tested, if it can be quantitatively demonstrated whether a new species looks very similar to the original taxon or whether its morphology has changed in a specific direction. For example, if a new species of herbivore evolves with increased grinding area in the cheek dentition, it can either occupy the same area of morphospace as previously existing species, suggesting increased resource competition, or it can occupy an area of morphospace that had previously been empty, suggesting evolution into a new niche. This example illustrates a situation where speciation did not just increase the number of taxa, but also morphologic and ecologic diversity. In turn, this quantitative information can be used to test speciation hypotheses in the extant fauna as well as the fossil record suggested by previous studies using molecular data and habitat reconstruction (Gaubert and Begg, 2007). Ecomorphology has been used at various scales to study the diversification of vertebrates (Van Valkenburgh, 1988, 1989; Werdelin, 1996; Wesley-Hunt, 2005), invertebrates (Foote, 1994, 1997), and plants, (Lupia, 1999) over their evolutionary history. It is akin to taxon-free analysis (Damuth, 1992), but whereas that mode of analysis seeks to use morphology to characterize paleoecological features of communities and habitats, the focus in ecomophology is on diversity of function within habitats or larger taxonomic groups. In this study, we apply ecomorphological analysis to the small Feliformia of the families Herpestidae and Viverridae. These taxa are all small tropical and subtropical carnivorans, distributionally limited to Africa and South and Southeast Asia (disregarding some introduced populations). In this paper we isolate these taxa, pulling them out of a much larger analysis including all modern terrestrial carnivorans (see Werdelin and Wesley-Hunt, chapter NN, this volume), to focus on their biogeographic and morphospace occupation patterns. The reason we analyze Herpestidae and Viverridae in this study is because of their shared ecology, ancestry, and biogeography. We have not included small Mustelidae, which share some similarities in ecology and morphology with Herpestidae and Viverridae, because although they may have some impact on the distribution patterns of herpestids and viverrids in Asia, in Africa they have not been a significant presence since the Early Miocene (Werdelin and Peigné, in press). Although competition with herpestids and viverrids (especially the former) may have influenced the representation of small Mustelidae in Africa, such interactions as took place 15-20 million years ago will have had little impact on the morphospace distributions of herpestids and viverrids in Africa today. Little is known about the evolutionary history and diversification of Herpestidae and Viverridae due to their generally poor fossil record. In addition, compared to the majority of Carnivora, little is known about the biology of their extant representatives due to the difficulties of studying them in the wild. Our aim here is to quantify their morphology and therefore some aspects of their ecology, in order to better understand the processes underlying their current distribution. The ability to distinguish dietary (ecological) categories has been demonstrated using dental and skull morphology in modern carnivorans (Radinsky, 1981a, b, 1982; VanValkenburgh, 1988, and more recently in using the small carnivoran dentition, Friscia et al. 2007). The detection of this pattern makes it possible to establish a set of morphological characters to infer the ecology of fossil taxa. Like many mammalian groups, Herpestidae and Viverridae have undergone extensive phylogenetic revision over the last decade, following the routine use of molecular methods in phylogenetic analysis. Most early authors placed these taxa in the single family Viverridae, with the majority of what are now viewed as Herpestidae placed in a subfamily Herpestinae (Gregory and Hellman, 1939). Later authors (e.g., Hunt, 1974) disagreed on morphological grounds with this placement, and with the introduction of molecular data, it was unequivocally shown that “Herpestinae” neither belonged within Viverridae, nor was it the sister taxon to that family, and the validity of a separate family Herpestidae was recognized (e.g., Flynn and Nedbal 1998). Our understanding of the relationships between species groups in the families Viverridae and Herpestidae has also changed drastically (Gaubert et al. 2002, 2004a, b, 2005; Veron et al. 2004; Gaubert and Cordeiro, 2006; Perez et al., 2006; Patou et al., 2008), and many subfamilies and genera are no longer considered monophyletic. Indeed, the Asiatic linsangs, hitherto considered to be Viverridae, have recently been shown to be the sister-group to the family Felidae (Gaubert and Veron, 2003). Further, it has been shown that the debated Malagasy carnivorans, earlier split among Viverridae and Herpestidae, and even Felidae, is a monophyletic sister-group to the Herpestidae (Veron and Catzeflis, 1993; Yoder et al., 2003; Flynn et al., 2005). These taxa are not included in the present analysis, which is concerned only with the monophyletic families Herpestidae and Viverridae, as currently conceived. The fossil record of both families is extremely sparse and difficult to interpret, compounding the confusion surrounding evolutionary history of these groups (Werdelin, 2003; Werdelin and Peigné, in press). Research continues to be done concerning these groups, but in the absence of a well-corroborated taxonomy and phylogeny, it has been very difficult to understand the determinants of their extensive radiation in the Old World. Ecomorphology uses morphological characters to assess the diversity of morphologies within a higher taxonomic unit. The target of analysis can be a clade, a guild, or a community, and the object is to link the morphology to the diversity of ecologies seen in the target group. The characters used will determine the type of ecological information included (feeding modes, locomotor categories, etc.). No study can address all aspects of an animal’s ecomorphology, and here we use characters that describe the dentition, coupled with body mass, the latter being a fundamental ecological parameter (Damuth and MacFadden, 1990; Owen-Smith, 1988). The characters were chosen to maximize the ecological and functional information available from macroscopic study of carnivoran dentition. This approach allows us to explore the radiation of Herpestidae and Viverridae in the absence of detailed knowledge of taxonomy and interrelationships. While our conclusions would be enhanced by better understanding of the evolutionary history of the taxa included, especially to examine if their morphospace occupation in deep time is constrained by their phylogeny, the present study is an important step toward teasing apart the evolutionary history and interconnectedness of these groups. METHODS The analysis on which this study is based was carried out on a set of 216 species, encompassing 85% of modern carnivoran species around the world. For each species, one specimen was selected to measure and code. This specimen was selected after studying a larger sample to ensure that the data collected represents an average individual, with no morphologic abnormalities. Due to the nature of the characters, which were designed to allow the extremes of Carnivora to be compared (a polar bear to a weasel); the variation among individuals of a species would only rarely result in a slight difference in the code, and a concomitant minute difference in the position in morphospace. From this study, the Herpestidae and Viverridae were extracted and their occupation of morphospace analyzed. The species studied include 27 Herpestidae and 23 Viverridae, representing approximately 82% and 80% respectively of existing species (Wozencraft, 1993) (Table 1). The specimens are housed in the following museums: Field Museum, Chicago; National Museum of Natural History, Smithsonian, Washington D.C.; American Museum of Natural History, New York; Museum für Naturkunde, Berlin; Museo
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