Abstract
Tooth root surface areas serve as proxies for bite force potentials, and by extension, dietary specialization in extant carnivorans. Here, we investigate the feeding ecology of the extinct large-bodied ursid Agriotherium africanum, by comparing its root surface areas (reconstructed with the aid of computed tomography and three-dimensional image processing) and bite force estimates, with those of extant carnivorans. Results show that in absolute terms, canine and carnassial bite forces, as well as root surface areas were highest in A. africanum. However, when adjusted for skull size, A. africanum’s canine roots were smaller than those of extant solitary predators. With teeth being the limiting factor in the masticatory system, low canine root surface areas suggest that A. africanum would have struggled to bring down large vertebrate prey. Its adjusted carnassial root sizes were found to be smaller than those of extant hard object feeders and the most carnivorous tough object feeders, but larger than those of extant omnivorous ursids and Ursus maritimus. This and the fact that it displayed its highest postcanine root surface areas in the carnassial region (rather than the most distal tooth in the tooth row) suggest that A. africanum consumed more vertebrate tissue than extant omnivorous ursids. With an apparent inability to routinely bring down large prey or to consume mechanically demanding skeletal elements, its focus was most likely on tough tissue, which it acquired by actively scavenging the carcasses of freshly dead/freshly killed animals. Mechanically less demanding skeletal elements would have been a secondary food source, ingested and processed mainly in association with muscle and connective tissue.
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References
Beeman LE, Pelton MR (1980) Seasonal foods and feeding ecology of black bears in the Smokey Mountains. Bears: Their Biology and Management 4:141–147
Bocherens H, Drucker DG, Billiou D, Geneste JM, van der Plicht J (2006) Bears and humans in Chauvet Cave (Vallon-Pont-d’Arc, Ardèche, France): insights from stable isotopes and radiocarbon dating of bone collagen. J Hum Evol 50:370–376
Bocherens H, Fizet M, Mariotti A (1990) Mise en evidence alimentaire vegetarien de l’ours des caverns (Ursus spelaeus) par la biogeochemie isotopique (13C, 15 N) des vertebres fossiles. Cr Acad Sci II 311:1279–1284
Bocherens H, Fizet M, Mariotti A (1994) Diet, physiology and ecology of fossil mammals as inferred from stable carbon and nitrogen isotope biogeochemistry: implications for Pleistocene bears. Palaeogeogr Palaeoclimatol Palaeoecol 107:213–225
Cane MA, Molnar P (2001) Closing of the Indonesian seaway as a precursor to East African aridification around 3–4 million years ago. Nature 411:157–162
Christiansen P (2007) Evolutionary implications of bite mechanics and feeding ecology in bears. J Zool London 272:423–443
Christiansen P (2008) Evolution of skull and mandible shape in cats (Carnivora: Felidae). Plos One 3(7): e2807
Christiansen P, Wroe S (2007) Bite forces and evolutionary adaptations to feeding ecology in carnivores. Ecology 88:347–358
Davis DD (1955) Masticatory apparatus in the spectacled bear Tremarctos ornatus. Fieldiana Zool 37:25–46
deMenocal PB (2004) African climate change and faunal evolution during the Pliocene-Pleistocene. Earth Planet Sci Lett 220:3–24
Derocher AE, Wiig Ø, Andersen M (2002) Diet composition of polar bears in Svalbard and the western Barents Sea. Polar Biol 25:448–452
Figueirido B, Soibelzon LH (2009) Inferring paleoecology in extinct tremarctine bears (Carnivora, Ursidae) via geometric morphometrics. Lethaia 43:209–222
Franz-Odendaal TA, Solounias N (2004) Comparative dietary evaluations of an extinct giraffid (Sivatherium hendeyi) (Mammalia, Giraffidae, Sivatheriinae) from Langebaanweg, South Africa (early Pliocene). Geodiversitas 26:675–685
Gentry A (1980) Fossil Bovidae (Mammalia) from Langebaanweg, South Africa. Ann S Afr Mus 79:213–337
Hammer Ø, Harper DAT, Ryan PD (2001) PAST: Paleontological statistics software package for education and data analysis. Palaeontol Electron 4: 9pp. http://palaeo-electronica.org/2001_1/past/issue1_01.htm
Hamon N, Emonet E-G, Chaimanee Y, Guy F, Tafforeau P, Jaeger JJ (2012) Analysis of dental root apical morphology: a new method for dietary reconstructions in primates. Anat Record doi:10.1002/ar.22482
Hendey QB (1972) A Pliocene ursid from South Africa. Ann S Afr Mus 59:115–132.
Hendey QB (1974) The late Cenozoic Carnivora of the south-western Cape Province. Ann S Afr Mus 63:1–369
Hendey QB (1977) Fossil bear from South Africa. S Afr J Sci 73:112–116
Hendey QB (1980) Agriotherium (Mammalia, Ursidae) from Langebaanweg, South Africa, and relationships of the genus. Ann S Afr Mus 81:1–109
Hobson KA, Stirling I (1997) Low variation in blood δ13C among Hudson Bay polar bears: implications for metabolism and tracing terrestrial foraging. Mar Mammal Sci 13:359–367
Jungers WL, Falsetti AB, Wall CE (1995) Shape, relative size, and size–adjustments in morphometrics. Am J Phys Anthropol 38:137–161
Klein RG (1982) Patterns of ungulate mortality and ungulate mortality profiles from Langebaanweg (early Pliocene) and Elandsfontein (middle Pleistocene), south-western Cape Province, South Africa. Ann S Afr Mus 90:49–94
Kovacs I (1971) A systematic description of dental roots. In: Dahlberg AA (ed) Dental Morphology and Evolution. University of Chicago Press, Chicago, pp 211–256
Kupczik K, Dean MC (2008) Comparative observations on the tooth root morphology of Gigantopithecus blacki. J Hum Evol 54:194–204
Kupczik K, Hublin JJ (2010) Mandibular molar root morphology in Neanderthals and late Pleistocene and recent Homo sapiens. J Hum Evol 59:525–541
Kupczik K, Stynder DD (2012) Tooth root morphology as an indicator for dietary specialization in carnivores (Mammalia: Carnivora). Biol J Linn Soc 105:456–471
Kurtén B (1967) Pleistocene bears of North America, II: Genus Arctodus, short-faced bears. Acta Zool Fenn 117:1–60
Lucas P, Constantino P, Wood B, Lawn B (2008) Dental enamel as a dietary indicator in mammals. Bioessays 30:374–385
Madić J, Huber D, Lugović B (1993) Serologic survey for selected viral and rickettsial agents of brown bears (Ursus arctos) in Croatia. J Wildlife Dis 29:572–576
Miles AEW, Grigson C (eds) (1990) Colyer’s Variations and Diseases of the Teeth of Animals. Cambridge University Press, Cambridge
Mowat G, Heard DC (2006) Major components of grizzly bear diet across North America. Can J Zool 84:473–489
Oldfield CC, McHenry CR, Clausen PD, Chamoli U, Parr WCH, Stynder DD, Wroe S (2012) Finite element analysis of ursid cranial mechanics and the prediction of feeding behaviour in the extinct giant Agriotherium africanum. J Zool London 286:93–171
Persson I-L, Wikan S, Swenson JE, Mysterud I (2001) The diet of the brown bear Ursus arctos in the Pasvik Valley, northeastern Norway. Wildlife Biol 7:27–37
Richards MP, Pacher M, Stiller M, Quilès J, Hofreiter M, Constantin S, Zilhao J, Trinkaus E (2008) Isotopic evidence for omnivory among European cave bears: late Pleistocene Ursus spelaeus from the Pestera cu Oase, Romania. Proc Natl Acad Sci USA 105:600–604
Sacco T, Van Valkenburgh B (2004) Ecomorphological indicators of feeding behaviour in the bears (Carnivora: Ursidae). J Zool London 263:41–54.
Schubert BW, Wallace SC (2009) Late Pleistocene giant shortfaced bears, mammoths, and large carcass scavenging in the Saltville Valley of Virginia, USA. Boreas 38:482–492
Sepulchre P, Ramstein G, Fluteau F, Schuster M, Tiercelin J-J, Brunet M (2006) Tectonic uplift and East African aridification. Science 313:1419–1423
Shrestha B, Reed JM, Starks PT, Kaufman GE, Goldstone JV, Roelke ME, O’Brien SJ, Koepfli K-P, Frank LG, Court MH (2011) Evolution of a major drug metabolizing enzyme defect in the domestic cat and other Felidae: phylogenetic timing and the role of hypercarnivory. Plos One 6: e18046
Slater GJ, Van Valkenburgh B (2008) Long in the tooth: evolution of sabertooth cat cranial shape. Paleobiology 34:403–419
Soibelzon LH, Schubert BW (2011) The largest known bear, Arctotherium angustidens, from the early Pleistocene Pampean region of Argentina: with a discussion of size and diet trends in bears. J Paleontol 85:69–75
Sorkin B (2006) Ecomorphology of the giant short-faced bears Agriotherium and Arctodus. Hist Biol 18:1–20
Spencer MA (1998) Force production in the primate masticatory system: electromyographic tests of biomechanical hypotheses. J Hum Evol 34:25–54
Spencer MA (2003) Tooth-root form and function in platyrrhine seed-eaters. Am J Phys Anthropol 122:325–335
Van Valkenburgh B (1989) Carnivore dental adaptations and diet: a study of trophic diversity within guilds. In: Gittleman JL (ed) Carnivore Behavior, Ecology, and Evolution. Chapman and Hall, London, pp 410–436
Van Valkenburgh B (2007) Déjà vu: the evolution of feeding morphologies in the Carnivora. Integr Comp Biol 47:147–163
Vinyard CJ, Wall CE, Williams SH, Hylander WL (2003) A comparative functional analysis of the skull morphology of tree gouging primates. Am J Phys Anthropol 120:153–170
Wroe S, Brammall J, Cooke B (1998) The skull of Ekaltadeta ima (Marsupialia, Hypsiprymnodontidae?): an analysis of some marsupial cranial features and a reinvestigation of propleopine phylogeny, with notes on the inference of carnivory in mammals. J Paleontol 72:738–751
Wroe S, McHenry C, Thomason JJ (2005) Bite club: comparative bite force in big biting mammals and the prediction of predatory behaviour in fossil taxa. Proc R Soc B-Biol Sci 272:619–625
Acknowledgements
We would like to thank P. Haarhoff (West Coast Fossil Park and Iziko South African Museum) for the loan of the A. africanum skull (SAM-PQL 45062) used in this study. N. Peters (Groote Schuur Hospital) is thanked for CT scanning assistance. The CT scanning of SAM-PQL 45062 was funded by a grant from the Palaeontological Scientific Trust (PAST) to D.D.S. and a National Research Foundation African Origins Platform Grant (AOP/West Coast Fossil Park) to R. Smith (Karoo Paleontology, Iziko SA Museum). We are grateful to Adam Sylvester for comments and advice on analysis. Comments from two anonymous reviewers significantly improved this manuscript.
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Stynder, D.D., Kupczik, K. Tooth Root Morphology in the Early Pliocene African Bear Agriotherium africanum (Mammalia, Carnivora, Ursidae) and its Implications for Feeding Ecology. J Mammal Evol 20, 227–237 (2013). https://doi.org/10.1007/s10914-012-9218-x
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DOI: https://doi.org/10.1007/s10914-012-9218-x