Chapter 7

Abstract

Borophaginae is the largest of the three subfamilies of the dog family Canidae, with some 66 species, spanning approximately 34 m.y. (Orellan to Blancan). Not surprisingly, this extensive radiation of canids includes a diverse array of dietary types, ranging from hypocarnivorous to hypercarnivorous and durophagous. The last 16 m.y. of borophagine history is dominated by hypercarnivorous forms that were the dominant doglike predators within their faunas. Because of their relatively robust skeletons and their resemblance to extant hyenas in craniodental morphology, many or most of these hypercarnivorous species, particularly those of the late Miocene and Pliocene, have been assumed as primarily scavengers rather than hunters. The classification of most hypercarnivorous borophagines as scavengers relegates them to much less important roles in the ecology and evolution of their respective communities than does a classification as hunters. Unlike hunters, scavengers are unlikely to influence the evolution of the animals they eat, and are expected to exist at relatively low densities as do the only extant scavenging carnivorans, brown (Parahyaena brunnea) and striped hyenas (Hyaena hyaena). Given the substantial fossil record of the Borophaginae, it seems unlikely that all or most of the hypercarnivorous forms were primarily scavengers. Moreover, if some hunted, the larger species might be expected to have done so in groups, as large canids hunt in packs today.Here we examine possible foraging modes within the Borophaginae using morphometrics and two new approaches to estimating the typical prey size of extinct carnivores. The craniodental morphology of the Borophaginae is compared with that of the living Caninae and Hyaeninae (hyaenids exclusive of Proteles cristata, the aardwolf) based on measurements that reflect relative tooth size, jaw muscle leverage, rigidity of the dentary, and grinding versus slicing function of the teeth. The Borophaginae are found to be intermediate in morphology between the Caninae and Hyaeninae. Unlike hyaenids and like canines, they retain substantial postcarnassial molars. However, like hyaenids, the borophagines had significantly stronger jaws and enhanced jaw muscle leverage compared to other canids. Prey size is estimated for borophagines based on correlation between dentary height and typical prey size in living canids. These results are compared with those produced using a recently published energetic model that predicts that all carnivores larger than about 21 kg feed on prey as large or larger than themselves. The methods provide similar predictions, resulting in a list of 11 borophagines (all subtribes Aelurodontina and Borophagina) that probably consumed large prey.Comparisons with extant hyaenids reveal that the sole hunter of large prey, the spotted hyena (Crocuta), differs from the two mainly scavenging species, the brown and striped hyenas, in being significantly larger, more abundant, and widespread. Moreover, morphometric comparisons indicate that spotted hyenas have a more hypercarnivorous dentition. Given this, it is expected that the largest, most common borophagines with the most reduced dental grinding areas hunted most of their food. Based on their craniodental morphology and abundance in the record, Epicyon saevus, E. haydeni, Borophagus secundus, Aelurodon ferox, and A. taxoides were hunters. Although it is clear that Aelurodon and Borophagus were more capable of grasping prey than are extant canids, no borophagine evolved sharp, curved claws as in felids. Consequently, their ability to grapple with prey seems to have been limited, and packs were probably more successful at making a kill than individuals. Previous workers have argued against hunting in borophagines based on heavy dental wear, robust skeletal morphology, and external brain features. None of these precludes either hunting or hunting in packs in our view, and sharp teeth are not required for making a kill. While li