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Linking Sexual Dimorphism and Sociality in Rhinoceroses: Insights from Teleoceras Proterum and Aphelops Malacorhinus from the Late Miocene of Florida

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Linking Sexual Dimorphism and Sociality in Rhinoceroses: Insights from Teleoceras Proterum and Aphelops Malacorhinus from the Late Miocene of Florida Bull. Fla. Mus. Nat. Hist. (2005) 45(4):495-520 495 LINKING SEXUAL DIMORPHISM AND SOCIALITY IN RHINOCEROSES: INSIGHTS FROM TELEOCERAS PROTERUM AND APHELOPS MALACORHINUS FROM THE LATE MIOCENE OF FLORIDA Matthew C. Mihlbachler1,2 A strong relationship between sexual dimorphism and the degree of polygyny (i.e., the degree to which males compete for mates) is not apparent in living perissodactyls. For instance, in both monomorphic and dimorphic species of rhinos, about half of male mortality is attributable to tusk and horn mediated combat. Males of the North American Miocene rhinoceros Teleoceras had delayed tusk (i2) eruption, prolonged tusk root growth, and highly sharpened tusk facets maintained by honing on the upper incisor, thus predicting high levels of intermale aggression similar to living rhinos. The sex biases and elevated male mortality rates found in Teleoceras assemblages from Nebraska and Florida seem to confirm this prediction. However, the degree of body size dimorphism in these assemblages varies. Therefore, the intensity of intermale competition seems unrelated to the magnitude of body size dimorphism in Teleoceras. Male individuals of Aphelops, a sympatric rhino, experienced more finite tusk growth and tusks were blunted with age due to the ancestral loss of the upper honing incisor, thus predicting lower levels of intermale competition. The Aphelops fossil assemblage from the Love Bone Bed of Florida is not sex-biased, shows more balanced sex-specific mortality rates, and seems to confirm the prediction of reduced intermale competition, thus suggesting a type of sociality that is different from both Teleoceras and modern rhinos. However, the same assemblage exhibits a degree of sexual dimorphism in tusk and body size that are is not demonstrably different from Teleoceras. Thus, we are left with a perplexing relationship between dimorphism and sociality for rhinos, where levels of intermale competition seem uncorrelated to the degree of sexual dimorphism in both living and extinct species. Key Words: Teleoceras; Aphelops; Miocene; rhinoceros; sexual dimorphism; sociality INTRODUCTION degree of polygyny in extinct species (Plavcan 2000). SEXUAL DIMORPHISM AND SOCIALITY IN PERISSODACTYLS The relationship of sexual dimorphism with sociality is Sexual dimorphism in mammals is best understood as a most clear among ruminant artiodactyls where increas- result of sexual selection in polygynous species where ingly larger species tend be extremely dimorphic and males invest large amounts of energy into competing form large, herd-like social groups that enable males to with other males to monopolize access to females monopolize large numbers of mates (Jarman 1983; Geist (Alexander et al. 1979; Jarman 1983, 2000; Clutton- & Bayer 1988; Loison et al 1999). Extant perissodac- Brock et al. 1988; Andersson 1994; Berger & tyls are unusual in the sense that, although all living spe- Cunningham 1994a). Mature males commonly possess cies appear to be polygynous, sexual dimorphism is not enlarged body size or enlarged weapon-like structures prominent and there is no clear-cut relationship between (e.g., horns, tusks, or antlers) that are used to compete sexual dimorphism and sociality. Tapirs and horses are for mates either directly through combat, or more indi- both monomorphic in body size, yet tapirs tend to be rectly through complex behaviors (e.g., ritualized dis- solitary while most horse species form year-round har- plays). The identification and quantification of sexual ems (Berger 1986; Rubenstein 1986; Nowak 1999). dimorphism in fossils is potentially informative of the Rhinos are the only living perissodactyls that posses con- spicuous weapon-like structures in the forms of horns 1 New York College of Osteopathic Medicine, Department of and tusks; therefore we might expect to find among rhi- Anatomy, Northern Boulevard, Old Westbury, NY 11568-8000. <[email protected]> nos a stronger relationship between dimorphism and so- 2 American Museum of Natural History, Division of Paleontol- ciality. The degree of sexual dimorphism in living rhinos ogy, Central Park West at 79th Street, New York, NY 10024 varies from species to species and there are both mono- 496 CENOZOIC VERTEBRATES: Papers to Honor S. David Webb morphic (Diceros bicornis) and moderately dimorphic the Ashfall site, Nebraska. Voorhies and Stover (1978) forms (Ceratotherium simum, Rhinoceros unicornis) found fetal bones within the abdominal regions of skel- (Groves 1982; Owen-Smith 1988; Dinerstein 1991, 1993; etons of small-tusked individuals from the Ashfall site, Berger 1994; Rachlow & Berger 1995). Despite some confirming that the small-tusked individuals were fe- variability in the degree of dimorphism, all living rhinos males. Matthew (1932) and Lambert (1994) found that are typically solitary. They rarely form small, tempo- lower tusks of Aphelops mutilus from Coffee Ranch, rary aggregations, and these appear to result from the Texas and Moss Acres, Florida could be easily divided chance clustering of critical food or water resources. into male and female groups based on size. Borsuk- Social bonding is minimal among adults, even among the Bialynicka (1973) discovered that several cranial dimen- most dimorphic species (Laurie 1982; Owen-Smith 1988; sions of Coelodonta antiquitatis, the Pleistocene woolly Dinerstein 2003). Despite the absence of coherent rhino of Europe, were bimodal. Finally, Deng (2001, groups, intermale competition among rhinos is intense 2005) attributed intraspecific variation in the cranial os- and they are among the most violent and aggressive of teologies of Chilotherium wimani and Iranotherium mammals. In the most well-studied populations of Af- morgani to sexual dimorphism. rica and Asia, aggressive confrontations among males Frequent sexual dimorphism among fossil perisso- account for up to half of male mortality in both mono- dactyls and less pronounced dimorphism among modern morphic (D. bicornis) and dimorphic species (C. simum species resembles Wright’s (1993) findings on peccar- and R. unicornis) (Hitchins & Anderson 1983; Owen- ies; sexual dimorphism is not prominent among the few Smith 1988; Dinerstein & Price 1991; Berger 1994; living species although pronounced dimorphism is found Berger & Cunningham 1994b; Dinerstein 2003). Males in extinct species. Perissodactyls were more diverse in are also aggressive towards females. For instance, R. the past, and extinction over the last several million years unicornis males ram females to subdue them (Dinerstein has resulted in drastic reductions in taxonomic diversity. 1991). Additionally, population sizes of most extant species have Sexual dimorphism is commonly reported for ex- been greatly diminished in recent times. Population tinct perissodactyls, suggesting that sexual dimorphism bottlenecking, geographic range restriction, artificial was more prominent in the past. Dimorphism has been population management, and phenotypic alteration (e.g., identified in an early “equoid” (Gingerich 1981), various dehorning of rhinos) can have dramatic effects on eco- ceratomorphs, including Homogalax, Isectolophus logical relationships and social behaviors of large mam- (Radinsky 1963), Hyrachyus (Radinsky 1967), mals (Berger 1994; Berger & Cunningham 1994a, Indricotherium transouralicum (Fortelius & 1994b). Consequently, recent perissodactyls might not Kappelman 1993), and the chalicothere Moropus serve as good models for the first ~99.9% of perisso- (Coombs 1975). Horned species of the extinct family dactyl evolutionary history. For instance, sociality and Brontotheriidae show high levels of intraspecific varia- sexual dimorphism might have been more strongly cor- tion in horn size and zygomatic thickness, suggesting a related in the past, during a time when anthropogenic degree of sexual dimorphism similar to modern rumi- effects were absent or minimal. nants (Osborn 1929; Mihlbachler et al. 2004a). Dimor- Among the three surviving families of perissodac- phism has been recognized among many extinct mem- tyls (Tapiridae, Equidae, Rhinocerotidae), sexual dimor- bers of the Rhinocerotidae. Osborn (1898a) reported phism appears to have been most pronounced in rhinos. dimorphism in the mandibular incisor and nasal horn The family Rhinocerotidae is cladistically defined by a bosses of Oligocene rhinos Subhyracodon occidentalis honing relationship between a chisel-like upper incisor and Diceratherium tridactylum. The early Miocene (I1) and a dimorphic tusk-like mandibular incisor (i2) Menoceras arikarense shows a degree of dimorphism (Prothero 2005). The living Asian rhinos (Rhinoceros in its horn bosses that is comparable to the level of di- unicornis, Rhinoceros sondaicus, and Dicerorhinus morphism found in modern ruminants (Peterson 1920; sumatrensis) possess both tusks and horns. Surpris- Mihlbachler unpublished data). Osborn (1898b) recog- ingly, the enlarged tusks, rather than the horns, are used nized sexual dimorphism in the tusks of Teleoceras to establish dominance hierarchies among Asian rhinos. fossiger from the late Miocene Long Island Rhino In R. unicornis, incisor size is strongly related to male Quarry, Kansas. Mead (2000) further quantified sexual dominance and tusk mediated combat commonly results dimorphism in tusk size and body size in Teleoceras from in mortal wounding (Laurie 1982; Dinerstein & Price MIHLBACHLER: Sexual Dimorphism and Sociality in Miocene Rhinoceroses 497 2001; Dinerstein 2003). African rhinos (Diceros this paper as well as results reported in Mihlbachler
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