Dogs: Their Fossil Relatives and Evolutionary History

FURTHER READING

THIS READING LIST OF PRIMARY LITERATURE for different topics is intended for serious readers to examine a particular subject further. It includes but also goes much beyond the literature cited in the text and provides a reasonably comprehensive list without delving too extensively into the details of historical treatments, such as living and fossil canid taxonomy.

ANATOMY, LOCOMOTION, AND FUNCTION

Andersson, K. 2004. Elbow-joint morphology as a guide to forearm function and foraging behaviour in mammalian carnivores. Zoological Journal of the Linnean Society 142:91–104.

Andersson, K., and L. Werdelin. 2003. The evolution of cursorial carnivores in the Tertiary: Implications of elbow-joint morphology. Biology Letters 270:163–165.

Antón, M., and A. Galobart 1999. Neck function and predatory behaviour in the scimitar tooth cat Homotherium latidens (Owen). Journal of Vertebrate Paleontology 19:771–784.

Baker, M. A., and L. W. Chapman. 1977. Rapid brain cooling in exercising dogs. Science 195:781–783.

Binder, W. J., and B. Van Valkenburgh. 2000. Development of bite strength and feeding behaviour in juvenile spotted hyenas (Crocuta crocuta). Journal of Zoology 252:273–283.

Carbone, C., G. M. Mace, S. C. Roberts, and D. W. Macdonald. 1999. Energetic constraints on the diet of terrestrial carnivores. Nature 402:286–288.

Carrano, M. T. 1997. Morphological indicators of foot posture in mammals: A statistical and biomechanical analysis. Zoological Journal of the Linnean Society 121:77–104.

Evans, H. E., and G. C. Christensen. 1979. Miller’s Anatomy of the Dog. Philadelphia: Saunders.

Ewer, R. F., and R. Singer. 1956. Fossil Carnivora from Hopefield. Annals of the South African Museum 42:335–347.

Flower, W. H. 1869. On the value of the characters of the base of the cranium in the classification of the order Carnivora, and on the systematic position of Bassaris and other disputed forms. Proceedings of the Zoological Society of London 1869:4–37.

Gaspard, M. 1964. La région de l’angle mandibulaire chez les Canidae. Mammalia 28:249–329.

Hildebrand, M. 1952. An analysis of body proportions in the Canidae. American Journal of Anatomy 90:217–256.

Hildebrand, M. 1954. Comparative morphology of the body skeleton in recent Canidae. University of California Publications in Zoology 52:399–470.

Holliday, J. A., and S. J. Steppan. 2004. Evolution of hypercarnivory: The effect of specialization on morphological and taxonomic diversity. Paleobiology 30:108–128.

Hunt, R. M., Jr. 2003. Intercontinental migration of large mammalian carnivores: Earliest occurrence of the Old World beardog Amphicyon (Carnivora, Amphicyonidae) in North America. Bulletin of the American Museum of Natural History 279:77–115.

Lee, D. V., J. E. A. Bertram, and R. J. Todhunter. 1999. Acceleration and balance in trotting dogs. Journal of Experimental Biology 202:3565–3573.

Munthe, K. 1989. The skeleton of the Borophaginae (Carnivora, Canidae): Morphology and function. University of California Publications in Geological Sciences 133:1–115.

Newman, C., C. D. Buesching, and J. O. Wolff. 2005. The function of facial masks in “midguild” carnivores. Oikos 108:623–633.

Ortolani, A. 1999. Spots, stripes, tail tips, and dark eyes: Predicting the function of carnivore colour patterns using the comparative method. Biological Journal of the Linnean Society 67:433–476.

Rensberger, J. M. 1995. Determination of stresses in mammalian dental enamel and their relevance to the interpretation of feeding behaviors in extinct taxa. In J. J. Thomason, ed., Functional Morphology in Vertebrate Paleontology, 151–172. Cambridge: Cambridge University Press.

Rensberger, J. M. 1997. Mechanical adaptation in enamel. In W. V. Koenigswald and P. M. Sander, eds., Tooth Enamel Microstructure, 237–257. Rotterdam: Balkema.

Stefen, C. 1999. Enamel microstructure of recent and fossil Canidae (Carnivora: Mammalia). Journal of Vertebrate Paleontology 19:576–587.

Usherwood, J. R., and A. M. Wilson. 2005. Biomechanics: No force limit on greyhound sprint speed. Nature 438:753.

Van Valkenburgh, B., and W. J. Binder. 2000. Biomechanics and feeding behavior in carnivores: Comparative and ontogenetic studies. In P. Domenici and R. W. Blake, eds., Biomechanics in Animal Bahaviour, 223–235. Oxford: BIOS.

Van Valkenburgh, B., and K.-P. Koepfli. 1993. Cranial and dental adaptations to predation in canids. In N. Dunstone and M. L. Gorman, eds., Mammals as Predators, 15–37. Oxford: Oxford University Press.

Van Valkenburgh, B., T. Sacco, and X. Wang. 2003. Pack hunting in Miocene borophagine dogs: Evidence from craniodental morphology and body size. Bulletin of the American Museum of Natural History 279:147–162.

Van Valkenburgh, B., J. Theodor, A. Friscia, A. Pollack, and T. Rowe. 2004. Respiratory turbinates of canids and felids: A quantitative comparison. Journal of Zoology 264:281–293.

Wang, X. 1993. Transformation from plantigrady to digitigrady: Functional morphology of locomotion in Hesperocyon (Canidae: Carnivora). American Museum Novitates 3069:1–23.

Wang, X., and B. M. Rothschild. 1992. Multiple hereditary osteochondromata of Oligocene Hesperocyon (Carnivora: Canidae). Journal of Vertebrate Paleontology 12:387–394.

Werdelin, L. 1989. Constraint and adaptation in the bone-cracking canid Osteoborus (Mammalia: Canidae). Paleobiology 15:387–401.

White, T. E. 1954. Preliminary analysis of the fossil vertebrates of the Canyon Ferry Reservoir area. Proceedings of the United States National Museum 103:395–438.

Wöhrmann-Repenning, A. 1993. The anatomy of the vermeronasal complex of the fox (Vulpes vulpes [L.]) under phylogenetic and functional aspects. Zoologische Jahrbucher: Abteilung für Anatomie und Ontogenie der Tiere 123:353–361.

Wroe, S., C. McHenry, and J. Thomason. 2005. Bite club: Comparative bite force in big biting mammals and the prediction of predatory behaviour in fossil taxa. Proceedings of the Royal Society of London, Series B 272:619–625.

BODY WEIGHT ESTIMATES

Andersson, K. 2004. Predicting carnivoran body mass from a weight-bearing joint. Journal of Zoology 262:161–172.

Anyonge, W., and C. Roman. 2006. New body mass estimates for Canis dirus, the extinct Pleistocene dire wolf. Journal of Vertebrate Paleontology 26:209–212.

Kaufman, J. A., and R. J. Smith. 2002. Statistical issues in the prediction of body mass for Pleistocene canids. Lethaia 35:32–34.

Van Valkenburgh, B. 1990. Skeletal and dental predictors of body mass in carnivores. In J. Damuth and B. MacFadden, eds., Body Size in Mammalian Paleobiology: Estimation and Biological Implications, 181–205. Cambridge: Cambridge University Press.

DOMESTICATION

Bardeleben, C., R. L. Moore, and R. K. Wayne. 2005. Isolation and molecular evolution of the Selenocysteine tRNA (Cf TRSP) and RNase P RNA (Cf RPPH1) genes in the dog family, Canidae. Molecular Biology and Evolution 22:347–359.

Cohn, J. 1997. How wild wolves became domestic dogs. BioScience 47:725–728.

Coppinger, R., and L. Coppinger. 2001. Dogs: A Startling New Understanding of Canine Origin, Behavior, and Evolution. New York: Scribner.

Davis, S. J. M., and F. R. Valla. 1978. Evidence for the domestication of the dog 12,000 years ago in the Natufian of Israel. Nature 276:608–610.

Dayan, T. 1994. Early domesticated dogs of the Near East. Journal of Archaeological Science 21:633–640.

Diamond, J. 1997. Guns, Germs, and Steel: The Fates of Human Societies. New York: Norton.

Hare, B., M. Brown, C. Williamson, and M. Tomasello. 2002. The domestication of social cognition in dogs. Science 298:1634–1636.

Hemmer, H. 1990. Domestication: The Decline of Environmental Appreciation. Cambridge: Cambridge University Press.

Leonard, J. A., R. K. Wayne, J. Wheeler, R. Valadez, S. Guillén, and C. Vilà. 2002. Ancient DNA evidence for Old World origin of New World dogs. Science 298:1613–1616.

Lindblad-Toh, K., C. M. Wade, T. S. Mikkelsen, E. K. Karlsson, D. B. Jaffe, M. Kamal, M. Clamp, J. L. Chang, E. J. Kulbokas, M. C. Zody, E. Mauceli, X. Xie, M. Breen, R. K. Wayne, E. A. Ostrander, C. P. Ponting, F. Galibert, D. R. Smith, P. J. deJong, E. Kirkness, P. Alvarez, T. Biagi, W. Brockman, J. Butler, C.-W. Chin, A. Cook, J. Cuff, M. J. Daly, D. DeCaprio, S. Gnerre, M. Grabherr, M. Kellis, M. Kleber, C. Bardeleben, L. Goodstadt, A. Heger, C. Hitte, L. Kim, K.-P. Koepfli, H. G. Parker, J. P. Pollinger, S. M. J. Searle, N. B. Sutter, R. Thomas, C. Webber, and E. S. Lander. 2005. Genome sequence, comparative analysis, and haplotype structure of the domestic dog. Nature 438:803–819.

Morey, D. E. 1994. The early evolution of the domestic dog. American Scientist 82:336–347.

Nobis, G. 1979. Der älteste Haushund lebte vor 14000 Jahren. Die Umschau 19:610.

Olsen, S. J. 1985. Origins of the Domestic Dog: The Fossil Record. Tucson: University of Arizona Press.

Olsen, S. J., and J. W. Olsen. 1977. The Chinese wolf, ancestor of New World dogs. Science 197:533–535.

Parker, H. G., L. V. Kim, N. B. Sutter, S. Carlson, T. D. Lorentzen, T. B. Malek, G. S. Johnson, H. B. DeFrance, E. A. Ostrander, and L. Kruglyak. 2004. Genetic structure of the purebred domestic dog. Science 304:1160–1164.

Sablin, M. V., and G. A. Khlopachev. 2002. The earliest Ice Age dogs: Evidence from Eliseevichi. Current Anthropology 43:795–799.

Savolainen, P., T. Leitner, A. N. Wilton, E. Matisoo-Smith, and J. Lundeberg. 2004. A detailed picture of the origin of the Australian dingo, obtained from the study of mitochondrial DNA. Proceedings of the National Academy of Sciences 101:12387–12390.

Savolainen, P., Y. Zhang, J. Luo, J. Lundeberg, and T. Leitner. 2002. Genetic evidence for an East Asian origin of domestic dogs. Science 298:1610–1613.

Schleidt, W. M., and M. D. Shalter. 2004. Co-evolution of humans and canids, an alternative view of dog domestication: Homo homini lupus? Evolution and Cognition 9:57–72.

Trut, L. N. 1999. Early canid domestication: The farm-fox experiment. American Scientist 87:160–169.

Vilà, C., P. Savolainen, J. E. Maldonado, I. R. Amorim, J. E. Rice, R. L. Honeycutt, K. A. Crandall, J. Lundeberg, and R. K. Wayne. 1997. The domestic dog has an ancient and genetically diverse origin. Science 276:1687–1689.

EVOLUTION

Baskin, J. A. 1998. Evolutionary trends in the late Miocene hyena-like dog Epicyon (Carnivora, Canidae). In Y. Tomida, L. J. Flynn, and L. J. Jacobs, eds., Advances in Vertebrate Paleontology and Geochronology, 191–214. Tokyo: National Science Museum.

Cope, E. D. 1880. Extinct Batrachia. American Naturalist 14:609–610.

Dayan, M., D. Simberloff, E. Tchernov, and Y. Yom-Tov. 1992. Canine carnassials: Character displacement in the wolves, jackals, and foxes of Israel. Biological Journal of the Linnean Society 45:315–331.

Dayan, M., E. Tchernov, Y. Yom-Tov, and D. Simberloff. 1989. Ecological character displacement in Saharo-Arabian Vulpes: Outfoxing Bergmann’s rule. Oikos 55:263–272.

Gittleman, J. L. 1986. Carnivore brain size, behavioral ecology, and phylogeny. Journal of Mammalogy 67:23–36.

Holliday, J. A., and S. J. Steppan. 2004. Evolution of hypercarnivory: The effect of specialization on morphological and taxonomic diversity. Paleobiology 30:108–128.

Kleiman, D. G., and J. F. Eisenberg. 1973. Comparisons of canid and felid social systems from an evolutionary perspective. Animal Behavior 21:637–659.

Muñoz-Durán, J. 2002. Correlates of speciation and extinction rates in the Carnivora. Evolutionary Ecology Research 4:963–991.

Radinsky, L. 1969. Outlines of canid and felid brain evolution. Annals of the New York Academy of Sciences 167:277–288.

Radinsky, L. 1973. Evolution of the canid brain. Brain Behavior and Evolution 7:169–202.

Turner, A., and M. Antón. 1996. The Big Cats and Their Fossil Relatives. New York: Columbia University Press.

Van Valkenburgh, B. 1991. Iterative evolution of hypercarnivory in canids (Mammalia: Carnivora): Evolutionary interactions among sympatric predators. Paleobiology 17:340–362.

Van Valkenburgh, B., and F. Hertel. 1993. Tough times at La Brea: Tooth breakage in large carnivores of the late Pleistocene. Science 261:456–459.

Werdelin, L., and M. E. Lewis. 2005. Plio-Pleistocene Carnivora of eastern Africa: Species richness and turnover patterns. Zoological Journal of the Linnean Society 144:121–144.

Wesley-Hunt, G. D., and J. J. Flynn. 2005. Phylogeny of the Carnivora: Basal relationships among the carnivoramorphans and assessment of the position of “Miacoidea” relative to Carnivora. Journal of Systematic Palaeontology 3:1–28.

FOSSIL SPECIES

Ballesio, R., and M. Philippe. 1995. Les Canidés Pléistocenes de la Balme a Collomb (commune d’Entremont-le-Vieux, Savoie). Nouvelles Archives du Muséum d’Histoire Naturelle de Lyon 33:45–65.

Barbour, E. H., and H. J. Cook. 1914. Two new fossil dogs of the genus Cynarctus from Nebraska. Nebraska Geological Survey 4:225–227.

Barbour, E. H., and H. J. Cook. 1917. Skull of Aelurodon platyrhinus sp. nov. Nebraska Geological Survey 7:173–180.

Barbour, E. H., and C. B. Schultz. 1935. A new Miocene dog, Mesocyon geringensis sp. nov. Bulletin of the Nebraska State Museum 1:407–418.

Baskin, J. A. 1980. The generic status of Aelurodon and Epicyon (Carnivora, Canidae). Journal of Paleontology 54:1349–1351.

Baskin, J. A. 2005. Carnivora from the late Miocene Love Bone Bed of Florida. Bulletin of the Florida Museum of Natural History 45:413–434.

Berry, C. T. 1938. A Miocene dog from Maryland. Proceedings of the United States National Museum 85:159–161.

Berta, A. 1981. Evolution of large canids in South America. Anais do II Congreso Latino-Americano de Paleontologia, Porto Alegre 2:835–845.

Berta, A. 1984. The Pleistocene bush dog Speothos pacivorus (Canidae) from the Lagoa Santa caves, Brazil. Journal of Mammalogy 65:549–559.

Berta, A. 1988. Quaternary evolution and biogeography of the large South American Canidae (Mammalia: Carnivora). University of California Publications in Geological Sciences 132:1–149.

Bever, G. S. 2005. Morphometric variation in the cranium, mandible, and dentition of Canis latrans and Canis lepophagus (Carnivora: Canidae) and its implications for the identification of isolated fossil specimens. Southwestern Naturalist 50:42–56.

Bjork, P. R. 1970. The Carnivora of the Hagerman Local Fauna (late Pliocene) of southwestern Idaho. Transactions of the American Philosophical Society, n.s., 60:1–54.

Bryant, H. N. 1991. Reidentification of the Chadronian supposed didelphid marsupial Alloeodectes mcgrewi as part of the deciduous dentition of the canid Hesperocyon. Canadian Journal of Earth Sciences 28:2062–2065.

Bryant, H. N. 1992. The Carnivora of the Lac Pelletier Lower Fauna (Eocene: Duchesnean), Cypress Hills Formation, Saskatchewan. Journal of Paleontology 66:847–855.

Bryant, H. N. 1993. Carnivora and Creodonta of the Calf Creek Local Fauna (late Eocene, Chadronian), Cypress Hills Formation, Saskatchewan. Journal of Paleontology 67:1032–1046.

Clark, J. 1939. Miacis gracilis, a new carnivore from the Unita Eocene (Utah). Annals of the Carnegie Museum 27:349–370.

Cook, H. J. 1909. Some new Carnivora from the lower Miocene beds of western Nebraska. Nebraska Geological Survey 3:262–272.

Cook, H. J. 1914. A new canid from the lower Pliocene of Nebraska. Nebraska Geological Survey 7:49–50.

Cook, H. J., and J. R. Macdonald. 1962. New Carnivora from the Miocene and Pliocene of western Nebraska. Journal of Paleontology 36:560–567.

Cope, E. D. 1873. Third notice of extinct Vertebrata from the Tertiary of the plains. Paleontological Bulletin 16:1–8.

Cope, E. D. 1877. Report upon the extinct Vertebrata obtained in New Mexico by parties of the expedition of 1874. In Report upon United States Geological Surveys West of the One Hundredth Meridian, First Lieut. Geo. M. Wheeler, Corps of Engineers, U.S. Army, in Charge, 4:1–370. Washington, D.C.: Government Printing Office.

Cope, E. D. 1883. On the extinct dogs of North America. American Naturalist 17:235–249.

Cope, E. D. 1884. The Vertebrata of the Tertiary formations of the West, book I. In Report upon United States Geological Surveys West of the One Hundredth Meridian, Part II, 3:1–1009. Washington, D.C.: Government Printing Office.

Cope, E. D. 1890. A new dog from the Loup Fork Miocene. American Naturalist 24:1067–1068.

Crusafont-Pairó, M. 1950. El primer representante del género Canis en el Pontiense eurasiatico (Canis cipio nova sp.). Boletín de la Real Sociedad Española de Historia Natural (Geología) 48:43–51.

Dayan, T. 1994. Carnivore diversity in the late Quaternary of Israel. Quaternary Research 41:343–349.

de Bonis, L., S. Peigné, A. Likius, H. T. Mackaye, P. Vignaud, and M. Brunet. 2007. The oldest African fox (Vulpes riffautae n. sp., Canidae, Carnivora) recovered in late Miocene deposits of the Djurab desert, Chad. Naturwissenschaften 94:575–580.

Emry, R. J., and R. E. Eshelman. 1998. The early Hemingfordian (early Miocene) Pollack Farm Local Fauna: First Tertiary land mammals described from Delaware. In R. N. Benson, ed., Geology and Paleontology of the Lower Miocene Pollack Farm Fossil Site, Delaware, 153–173. Delaware Geological Survey Special Publication no. 21. Newark: Delaware Geological Survey.

Evander, R. L. 1986. Carnivores of the Railway Quarries Local Fauna. Transactions of the Nebraska Academy of Science 14:25–34.

Eyerman, J. 1894. Preliminary notice of a new species of Temnocyon and a new genus from the John Day Miocene of Oregon. American Geologist 14:320–321.

Eyerman, J. 1896. The genus Temnocyon and a new species thereof and the new genus Hypotemnodon, from the John Day Miocene of Oregon. American Geologist 17:267–286.

Frailey, D. 1978. An early Miocene (Arikareean) fauna from north central Florida (the SB-1A Local Fauna). Occasional Papers of the Museum of Natural History, University of Kansas 75:1–20.

Frailey, D. 1979. The large mammals of the Buda Local Fauna (Arikareean: Alachua County, Florida). Bulletin of the Florida State Museum, Biological Sciences 24:123–173.

Galbreath, E. C. 1953. A contribution to the Tertiary geology and paleontology of northeastern Colorado. University of Kansas Paleontological Contributions 4:1–120.

Galbreath, E. C. 1956. Remarks on Cynarctoides arcidens from the Miocene of northeastern Colorado. Transactions of the Kansas Academy of Science 59:373–378.

Gawne, C. E. 1975. Rodents from the Zia Sand Miocene of New Mexico. American Museum Novitates 2586:1–25.

Gazin, C. L. 1932. A Miocene mammalian fauna from southeastern Oregon. Carnegie Institution of Washington, Contributions to Paleontology 418:37–86.

Gazin, C. L. 1942. The late Cenozoic vertebrate faunas from the San Pedro Valley, Ariz. Proceedings of the United States National Museum 92:475–518.

Geraads, D. 1997. Carnivores du Pliocène terminal de Ahl al Oughlam (Casablanca, Maroc). Geobios 30:127–164.

Green, M. 1948. A new species of dog from the lower Pliocene of California. University of California Publications, Bulletin of the Department of Geological Sciences 28:81–90.

Green, M. 1954. A cynarctine from the upper Oligocene of South Dakota. Transactions of the Kansas Academy of Science 57:218–220.

Gustafson, E. P. 1986. Carnivorous mammals of the late Eocene and early Oligocene of Trans-Pecos Texas. Bulletin of the Texas Memorial Museum 33:1–66.

Hall, E. R., and W. W. Dalquest. 1962. A new doglike carnivore, genus Cynarctus, from the Clarendonian, Pliocene, of Texas. University of Kansas Publications, Museum of Natural History 14:137–138.

Harrison, J. A. 1983. The Carnivora of the Edson Local Fauna (late Hemphillian), Kansas. Smithsonian Contributions to Paleobiology 54:1–42.

Hayes, F. G. 2000. The Brooksville 2 Local Fauna (Arikareean, latest Oligocene): Hernando County, Florida. Bulletin of the Florida Museum of Natural History 43:1–47.

Henshaw, P. C. 1942. A Tertiary mammalian fauna from the San Antonio Mountains near Tonopah, Nevada. Carnegie Institution of Washington, Contributions to Paleontology 530:77–168.

Hesse, C. J. 1936. A Pliocene vertebrate fauna from Optima, Oklahoma. University of California Publications, Bulletin of the Department of Geological Sciences 24:57–70.

Hibbard, C. W. 1950. Mammals of the Rexroad Formation from Fox Canyon, Meade County, Kansas. Contributions from the Museum of Paleontology, University of Michigan 8:113–192.

Hough, J. R., and R. Alf. 1956. A Chadron mammalian fauna from Nebraska. Journal of Paleontology 30:132–140.

Johnston, C. S. 1937. Tracks from the Pliocene of west Texas. American Midland Naturalist 28:147–152.

Johnston, C. S. 1939. A skull of Osteoborus validus from the early middle Pliocene of Texas. Journal of Paleontology 13:526–530.

Koufos, G. D. 1992. The Pleistocene carnivores of the Mygdonia basin (Macedonia, Greece). Annales de Paléontologie 78:205–257.

Koufos, G. D. 1997. The canids Eucyon and Nyctereutes from the Ruscinian of Macedonia, Greece. Paleontologia i Evolució 30–31:39–48.

Kurtén, B. 1974. A history of coyote-like dogs (Canidae, Mammalia). Acta Zoologica Fennica 140:1–38.

Kurtén, B. 1984. Geographic differentiation in the Rancholabrean dire wolf (Canis dirus Leidy) in North America. In H. H. Genoways and M. R. Dawson, eds., Contributions in Quaternary Vertebrate Paleontology: A Volume in Memorial to John E. Guilday, 218–227. Pittsburgh: Carnegie Museum of Natural History.

Leidy, J. 1858. Notice of remains of extinct Vertebrata, from the valley of the Niobrara River, collected during the exploring expedition of 1857, in Nebraska, under the command of Lieut. G. K. Warren, U.S. Top. Eng., by Dr. F. V. Hayden, geologist to the expedition. Proceedings of the Academy of Natural Sciences of Philadelphia 1858:20–29.

Leidy, J. 1869. The extinct mammalian fauna of Dakota and Nebraska, including an account of some allied forms from other localities, together with a synopsis of the mammalian remains of North America. Journal of the Academy of Natural Sciences of Philadelphia 7:1–472.

Loomis, F. B. 1931. A new Oligocene dog. American Journal of Science 22:100–102.

Loomis, F. B. 1932. The small carnivores of the Miocene. American Journal of Science 24:316–329.

Loomis, F. B. 1936. Three new Miocene dogs and their phylogeny. Journal of Paleontology 10:44–52.

Lyras, G. A., A. A. E. Van der Geer, M. D. Dermitzakis, and J. D. Vos. 2006. Cynotherium sardous, an insular canid (Mammalia: Carnivora) from the Pleistocene of Sardinia (Italy), and its origin. Journal of Vertebrate Paleontology 28:735–745.

Macdonald, J. R. 1948. The Pliocene carnivores of the Black Hawk Ranch Fauna. University of California Publications, Bulletin of the Department of Geological Sciences 28:53–80.

Macdonald, J. R. 1963. The Miocene faunas from the Wounded Knee area of western South Dakota. Bulletin of the American Museum of Natural History 125:141–238.

Macdonald, J. R. 1967a. A new species of late Oligocene dog, Brachyrhynchocyon sesnoni, from South Dakota. Contributions in Science, Los Angeles County Museum 126:1–5.

Macdonald, J. R. 1967b. A new species of late Oligocene dog, Sunkahetanka sheffleri, from South Dakota. Contributions in Science, Los Angeles County Museum 127:1–5.

Macdonald, J. R. 1970. Review of the Miocene Wounded Knee faunas of southwestern South Dakota. Bulletin of the Los Angeles County Museum of Natural History 8:1–82.

Martin, H. T. 1928. Two new carnivores from the Pliocene of Kansas. Journal of Mammalogy 9:233–236.

Martin, R. 1971. Les affinités de Nyctereutes megamastoides (Pomel) canidé du gisement Villafranchien de Saint-Vallier (Drôme, France). Palaeovertebrata 4:39–58.

Martin, R. 1973. Trois nouvelles espèces de Caninae (Canidae, Carnivora) des gisements plio-villafranchiens d’Europe. Documents des Laboratoires de Géologie de la Faculté des Sciences de Lyon 57:87–96.

Matthew, W. D. 1901. Fossil mammals of the Tertiary of northeastern Colorado. Memoirs of the American Museum of Natural History 1:355–448.

Matthew, W. D. 1902. New Canidae from the Miocene of Colorado. Bulletin of the American Museum of Natural History 16:281–290.

Matthew, W. D. 1918. Contributions to the Snake Creek fauna with notes upon the Pleistocene of western Nebraska, American Museum Expedition of 1916. Bulletin of the American Museum of Natural History 38:183–229.

Matthew, W. D. 1924. Third contribution to the Snake Creek fauna. Bulletin of the American Museum of Natural History 50:59–210.

McGrew, P. O. 1935. A new Cynodesmus from the lower Pliocene of Nebraska with notes on the phylogeny of dogs. University of California Publications, Bulletin of the Department of Geological Sciences 23:305–312.

McGrew, P. O. 1937. The genus Cynarctus. Journal of Paleontology 11:444–449.

McGrew, P. O. 1938. Dental morphology of the Procyonidae with a description of Cynarctoides, gen. nov. Field Museum of Natural History, Geological Series 6:323–339.

McGrew, P. O. 1941. A new procyonid from the Miocene of Nebraska. Field Museum of Natural History, Geological Series 8:33–36.

McGrew, P. O. 1944a. The Aelurodon saevus group. Field Museum of Natural History, Geological Series 8:79–84.

McGrew, P. O. 1944b. An Osteoborus from Honduras. Field Museum of Natural History, Geological Series 8:75–77.

McKenna, M. C., and S. K. Bell. 1997. Classification of Mammals Above the Species Level. New York: Columbia University Press.

Merriam, J. C. 1903. The Pliocene and Quaternary Canidae of the Great Valley of California. University of California Publications, Bulletin of the Department of Geological Sciences 3:277–290.

Merriam, J. C. 1906. Carnivora from the Tertiary formations of the John Day region. University of California Publications, Bulletin of the Department of Geological Sciences 5:1–64.

Merriam, J. C. 1911. Tertiary mammal beds of Virgin Valley and Thousand Creek in northwestern Nevada. University of California Publications, Bulletin of the Department of Geological Sciences 6:199–306.

Merriam, J. C. 1913. Notes on the canid genus Tephrocyon. University of California Publications, Bulletin of the Department of Geological Sciences 7:359–372.

Merriam, J. C. 1919. Tertiary mammalian faunas of the Mohave Desert. University of California Publications, Bulletin of the Department of Biological Sciences 11:437–585.

Merriam, J. C., and W. J. Sinclair. 1907. Tertiary faunas of the John Day region. University of California Publications, Bulletin of the Department of Geological Sciences 5:171–205.

Miller, W. E., and O. Carranza-Castañeda. 1998. Late Tertiary canids from central Mexico. Journal of Paleontology 72:546–556.

Morales, J., M. Pickford, and D. Soria. 2005. Carnivores from the late Miocene and basal Pliocene of the Tugen Hills, Kenya. Revista de la Sociedad Geológica de España 18:39–61.

Munthe, K. 1998. Canidae. In C. M. Janis, K. M. Scott, and L. L. Jacobs, eds., Evolution of Tertiary Mammals of North America, vol. 1, Terrestrial Carnivores, Ungulates, and Ungulatelike Mammals, 124–143. Cambridge: Cambridge University Press.

Nowak, R. M. 1979. North American Quaternary Canis. Monograph of the Museum of Natural History, University of Kansas 6:1–154.

Obara, I., and Y. Hasegawa. 2003. A skull of the Japanese wolf, Canis hodophilax Temminck, found in Ogura-yama limestone fissure, Ueno-mura, Gunma Prefecture. Bulletin of the Gunma Museum of Natural History 7:35–39.

Olsen, S. J. 1956a. The Caninae of the Thomas Farm Miocene. Breviora 26:1–12.

Olsen, S. J. 1956b. A new species of Osteoborus from the Bone Valley Formation of Florida. Florida Geological Survey Special Publication 2:1–5.

Olson, E. C., and P. O. McGrew. 1941. Mammalian fauna from the Pliocene of Honduras. Bulletin of the Geological Society of America 52:1219–1244.

Peterson, O. A. 1910. Description of new carnivores from the Miocene of western Nebraska. Memoirs of the Carnegie Museum 4:205–278.

Peterson, O. A. 1924. Discovery of fossil mammals in the Brown’s Park Formation of Moffatt County, Colorado. Annals of the Carnegie Museum 15:299–304.

Prevosti, F. J., A. E. Zuritaa, and A. A. Carlini. 2005. Biostratigraphy, systematics, and paleoecology of Protocyon Giebel, 1855 (Carnivora, Canidae) in South America. Journal of South American Earth Sciences 20:5–12.

Qiu, Z., and R. H. Tedford. 1990. A Pliocene species of Vulpes from Yushe, Shanxi. Vertebrata PalAsiatica 28:245–258.

Richey, K. A. 1938. Osteoborus diabloensis, a new dog from the Black Hawk Ranch fauna, Mount Diablo, California. University of California Publications, Bulletin of the Department of Geological Sciences 24:303–307.

Richey, K. A. 1979. Variation and evolution in the premolar teeth of Osteoborus and Borophagus (Canidae). Transactions of the Nebraska Academy of Science 7:105–123.

Riggs, E. S. 1942. Preliminary description of two lower Miocene carnivores. Field Museum of Natural History, Geological Series 7:59–62.

Romer, A. S., and A. H. Sutton. 1927. A new arctoid carnivore from the lower Miocene. American Journal of Science 14:459–464.

Rook, L. 1992. “Canis” monticinensis sp. nov., a new Canidae (Carnivora, Mammalia) from the late Messinian of Italy. Bolletino della Società Paleontologica Italiana 31:151–156.

Rook, L. 1994. The Plio-Pleistocene Old World Canis (Xenocyon) ex gr. falconeri. Bolletino della Società Paleontologica Italiana 33:71–82.

Rook, L., and D. Torre. 1996a. The latest Villafranchian–early Galerian small dogs of the Mediterranean area. Acta Zoologica Cracoviensia 39:427–434.

Rook, L., and D. Torre. 1996b. The wolf-event in western Europe and the beginning of the late Villafranchian. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 1996:495–501.

Russell, L. S. 1934. Revision of the lower Oligocene vertebrate fauna of the Cypress Hills, Saskatchewan. Transactions of the Royal Canadian Institute 20:49–67.

Russell, R. D., and V. L. VanderHoof. 1931. A vertebrate fauna from a new Pliocene formation in northern California. University of California Publications, Bulletin of the Department of Geological Sciences 20:11–21.

Savage, D. E. 1941. Two new middle Pliocene carnivores from Oklahoma, with notes on the Optima fauna. American Midland Naturalist 25:692–710.

Schlaikjer, E. M. 1935. Contributions to the stratigraphy and paleontology of the Goshen Hole area, Wyoming. IV. New vertebrates and the stratigraphy of the Oligocene and early Miocene. Bulletin of the Museum of Comparative Zoology 76:97–189.

Scott, W. B. 1890a. The dogs of the American Miocene. Princeton College Bulletin 2:37–39.

Scott, W. B. 1890b. Preliminary account of the fossil mammals from the White River and Loup Fork formations, contained in the Museum of Comparative Zoology. Pt. II. The Carnivora and Artiodactyla. Bulletin of the Museum of Comparative Zoology 20:65–100.

Scott, W. B. 1893. The mammals of the Deep River beds. American Naturalist 27:659–662.

Scott, W. B. 1897. Preliminary notes on the White River Canidae. Princeton University Bulletin 9:1–3.

Scott, W. B. 1898. Notes on the Canidae of the White River Oligocene. Transactions of the American Philosophical Society 19:325–415.

Scott, W. B., and G. L. Jepsen. 1936. The mammalian fauna of the White River Oligocene. Princeton University Bulletin 28:1–980.

Sellards, E. H. 1916. Fossil vertebrates from Florida: A new Miocene fauna; new Pliocene species; the Pleistocene fauna. Florida State Geological Survey Annual Report 8:77–160.

Simpson, G. G. 1930. Tertiary land mammals of Florida. Bulletin of the American Museum of Natural History 59:149–211.

Simpson, G. G. 1932. Miocene land mammals from Florida. Bulletin of the Florida State Geological Survey 10:1–41.

Sinclair, W. J. 1915. Additions to the fauna of the lower Pliocene Snake Creek beds (results of the Princeton University 1914 expedition to Nebraska). Proceedings of the American Philosophical Society 54:73–95.

Soria, D., and E. Aguirre. 1976. El cánido de Layna: Revisión de los Nyctereutes fósiles. In M. T. Alberdi and E. Aguirre, eds., Miscelanea neogena, 83–116. Madrid: International Geological Correlation Program.

Sotnikova, M. V. 2001. Remains of Canidae from the lower Pleistocene site of Untermassfeld. In R.-D. Kahlke, ed., Das Pleistozäne von Untermassfeld bei Meiningen (Thüringgen), pt. 2, 607–632. Mainz: Römisch-Germanischen Zentralmuseums.

Sotnikova, M.V. 2006. A new canid Nurocyon chonokhariensis gen. et sp. nov. (Canini, Canidae, Mammalia) from the Pliocene of Mongolia. Courier Forschungsinstitut Senckenberg 256:11–21.

Stevens, M. S. 1977. Further study of Castolon Local Fauna (early Miocene), Big Bend National Park, Texas. Pearce-Sellards Series of the Texas Memorial Museum 28:1–69.

Stevens, M. S. 1991. Osteology, systematics, and relationships of earliest Miocene Mesocyon venator (Carnivora: Canidae). Journal of Vertebrate Paleontology 11:45–66.

Stiner, M. C., F. C. Howell, B. Martinez-Navarro, E. Tchernov, and O. Bar-Yosef. 2001. Outside Africa: Middle Pleistocene Lycaon from Hayonim Cave, Israel. Bolletino della Società Paleontologica Italiana 40:293–302.

Stirton, R. A., and V. L. VanderHoof. 1933. Osteoborus, a new genus of dogs, and its relation to Borophagus Cope. University of California Publications, Bulletin of the Department of Geological Sciences 23:175–182.

Stock, C. 1928. Canid and proboscidian remains from the Ricardo deposits, Mohave Desert, California. Carnegie Institution of Washington, Contributions to Paleontology 393:39–49.

Stock, C. 1932. Hyaenognathus from the late Pliocene of the Coso Mountain, California. Journal of Mammalogy 13:263–266.

Stock, C. 1933. Carnivora from the Sespe of the Las Posas Hills, California. Carnegie Institution of Washington, Contributions to Paleontology 440:29–42.

Stock, C., and E. L. Furlong. 1926. New canid and rhinocerotid remains from the Ricardo Pliocene of the Mojave Desert, California. University of California Publications, Bulletin of the Department of Biological Sciences 16:43–60.

Tanner, L. G. 1973. Notes regarding skull characteristics of Oxetocyon cuspidatus Green (Mammalia, Canidae). Transactions of the Nebraska Academy of Science 2:66–69.

Tedford, R. H. 1978. History of dogs and cats: A view from the fossil record. In Nutrition and Management of Dogs and Cats, chap. M23. St. Louis: Ralston Purina.

Tedford, R. H. 1981. Mammalian biochronology of the late Cenozoic basins of New Mexico. Bulletin of the Geological Society of America 91:1008–1022.

Tedford, R. H., and D. Frailey. 1976. Review of some Carnivora (Mammalia) from the Thomas Farm local fauna (Hemingfordian: Gilchrist County, Florida). American Museum Novitates 2610:1–9.

Tedford, R. H., and Z. Qiu. 1991. Pliocene Nyctereutes (Carnivora: Canidae) from Yushe, Shanxi, with comments on Chinese fossil raccoon-dogs. Vertebrata PalAsiatica 29:176–189.

Tedford, R. H., and Z. Qiu. 1996. A new canid genus from the Pliocene of Yushe, Shanxi Province. Vertebrata PalAsiatica 34:27–40.

Tedford, R. H., and X. Wang. 2008. Metalopex, a new genus of fox (Carnivora: Canidae: Vulpini) from the late Miocene of western North America. Contributions in Science, Natural History Museum of Los Angeles County 41:273–278.

Tedford, R. H., X. Wang, and B. E. Taylor. In press. Phylogenetic systematics of the North American fossil Caninae (Carnivora: Canidae). Bulletin of the American Museum of Natural History.

Thorpe, M. R. 1922a. Oregon Tertiary Canidae, with descriptions of new forms. American Journal of Science 3:162–176.

Thorpe, M. R. 1922b. Some Tertiary Carnivora in the Marsh Collection, with descriptions of new forms. American Journal of Science 3:423–455.

VanderHoof, V. L. 1931. Borophagus littoralis from the marine Tertiary of California. University of California Publications, Bulletin of the Department of Geological Sciences 21:15–24.

VanderHoof, V. L. 1936. Notes on the type Borophagus diversidens Cope. Journal of Mammalogy 17:415–516.

VanderHoof, V. L., and J. T. Gregory. 1940. A review of the genus Aelurodon. University of California Publications, Bulletin of the Department of Geological Sciences 25:143–164.

Voorhies, M. R. 1965. The Carnivora of the Trail Creek Fauna. Contributions to Geology, University of Wyoming 4:21–25.

Wang, X. 1990. Pleistocene dire wolf remains from the Kansas River with notes on dire wolves in Kansas. Occasional Papers of the Museum of Natural History, University of Kansas 137:1–7.

Wang, X. 1994. Phylogenetic systematics of the Hesperocyoninae (Carnivora: Canidae). Bulletin of the American Museum of Natural History 221:1–207.

Wang, X. 2003. New material of Osbornodon from the early Hemingfordian of Nebraska and Florida. Bulletin of the American Museum of Natural History 279:163–176.

Wang, X., and R. H. Tedford. 1992. The status of genus Nothocyon Matthew, 1899 (Carnivora): An arctoid not a canid. Journal of Vertebrate Paleontology 12:223–229.

Wang, X., and R. H. Tedford. 1996. Canidae. In D. R. Prothero and R. J. Emry, eds., The Terrestrial Eocene–Oligocene Transition in North America, pt. 2, Common Vertebrates of the White River Chronofauna, 433–452. Cambridge: Cambridge University Press.

Wang, X., and R. H. Tedford. 2007. Evolutionary history of canids. In P. Jensen, ed., The Behavioural Biology of Dogs, 3–20. Oxford: CABI International.

Wang, X., and R. H. Tedford. 2008. Fossil dogs (Carnivora, Canidae) from the Sespe and Vaqueros formations in southern California, with comments on relationships of Phlaocyon taylori. Contributions in Science, Natural History Museum of Los Angeles County 41:255–272.

Wang, X., R. H. Tedford, and B. E. Taylor. 1999. Phylogenetic systematics of the Borophaginae (Carnivora: Canidae). Bulletin of the American Museum of Natural History 243:1–391.

Wang, X., R. H. Tedford, B. Van Valkenburgh, and R. K. Wayne. 2004a. Ancestry: Evolutionary history, molecular systematics, and evolutionary ecology of Canidae. In D. W. Macdonald and C. Sillero-Zubiri, eds., The Biology and Conservation of Wild Canids, 39–54. Oxford: Oxford University Press.

Wang, X., R. H. Tedford, B. Van Valkenburgh, and R. K. Wayne. 2004b. Phylogeny, classification, and evolutionary ecology of the Canidae. In C. Sillero-Zubiri, M. Hoffmann, and D. W. Macdonald, eds., Canids: Foxes, Wolves, Jackals, and Dogs. Status Survey and Conservation Action Plan, 8–20. Gland, Switzerland: International Union for the Conservation of Nature and Natural Resources, Species Survival Commission, Canid Specialist Group, World Conservation Union.

Wang, X., B. C. Wideman, R. Nichols, and D. L. Hanneman. 2004. A new species of Aelurodon (Carnivora, Canidae) from the Barstovian of Montana. Journal of Vertebrate Paleontology 24:445–452.

Webb, S. D. 1969a. The Burge and Minnechaduza Clarendonian mammalian fauna of north-central Nebraska. University of California Publications in Geological Sciences 78:1–191.

Webb, S. D. 1969b. The Pliocene Canidae of Florida. Bulletin of the Florida State Museum, Biological Sciences 14:273–308.

Webb, S. D., B. J. MacFadden, and J. A. Baskin. 1981. Geology and paleontology of the Love Bone bed from the late Miocene of Florida. American Journal of Science 281:513–544.

Webb, S. D., and S. C. Perrigo. 1984. Late Cenozoic vertebrates from Honduras and El Salvador. Journal of Vertebrate Paleontology 4:237–254.

Werdelin, L., and M. E. Lewis. 2005. Plio-Pleistocene Carnivora of eastern Africa: Species richness and turnover patterns. Zoological Journal of the Linnean Society 144:121–144.

White, T. E. 1941. Additions to the fauna of the Florida Pliocene. Proceedings of the New England Zoological Club 18:67–70.

White, T. E. 1942. The lower Miocene mammal fauna of Florida. Bulletin of the Museum of Comparative Zoology 92:1–49.

White, T. E. 1947. Addition to the Miocene fauna of north Florida. Bulletin of the Museum of Comparative Zoology 99:497–515.

Wilson, J. A. 1939. A new species of dog from the Miocene of Colorado. Contributions from the Museum of Paleontology, University of Michigan 5:315–318.

Wilson, J. A. 1960. Miocene carnivores, Texas coastal plain. Journal of Paleontology 34:983–1000.

Wortman, J. L., and W. D. Matthew. 1899. The ancestry of certain members of the Canidae, the Viverridae, and Procyonidae. Bulletin of the American Museum of Natural History 12:109–139.

MODERN SPECIES

Ansorge, H. 1994. Intrapopula skull variability in the red fox, Vulpes vulpes (Mammalia: Carnivora: Canidae). Zoologische Abhandlungen 48:103–123.

Audet, A. M., C. B. Robbins, and S. Larivière. 2002. Alopex lagopus. Mammalian Species 713:1–10.

Beisiegel, B. D. M., and G. L. Zuercher. 2005. Speothos venaticus. Mammalian Species 783:1–6.

Bekoff, M. 1977. Canis latrans. Mammalian Species 79:1–9.

Berta, A. 1982. Cerdocyon thous. Mammalian Species 186:1–4.

Berta, A. 1986. Aletocynus microtis. Mammalian Species 256:1–3.

Cabrera, A. 1931. On some South America canine genera. Journal of Mammalogy 12:54–67.

Churcher, C. S. 1960. Cranial variation in the North American red fox. Journal of Mammalogy 41:349–360.

Clark, H. O. 2005. Otocyon megalotis. Mammalian Species 766:1–5.

Cohen, J. A. 1978. Cuon alpinus. Mammalian Species 100:1–3.

Dietz, J. M. 1985. Chrysocyon brachyurus. Mammalian Species 234:1–4.

Egoscue, H. J. 1979. Vulpes velox. Mammalian Species 122:1–5.

Frafjord, K., and I. Stevy. 1998. The red fox in Norway: Morphological adaptation or random variation in size? Zeitschrift für Säugetierkunde 63:16–25.

Fritzell, E. K., and K. J. Haroldson. 1982. Urocyon cinereoargenteus. Mammalian Species 198:1–8.

Fuller, T. K., and P. W. Kat. 1993. Hunting success of African wild dogs in southwestern Kenya. Journal of Mammalogy 74:464–467.

Fuller, T. K., T. H. Nicholls, and P. W. Kat. 1995. Prey and estimated food consumption of African wild dogs in Kenya. South African Journal of Wildlife Research 25:3.

Geffen, E., M. E. Gompper, J. L. Gittleman, H.-K. Luh, D. W. Macdonald, and R. K. Wayne. 1996. Size, life-history traits, and social organization in the Canidae: A reevaluation. American Naturalist 147:140–160.

Geffen, E., R. Hefner, D. W. Macdonald, and M. Ucko. 1992. Diet and foraging behavior of Blanford’s foxes, Vulpes cana, in Israel. Journal of Mammalogy 73:395–402.

Gittleman, J. L. 1986. Carnivore brain size, behavioral ecology, and phylogeny. Journal of Mammalogy 67:23–36.

Gittleman, J. L., and S. L. Pimm. 1991. Crying wolf in North America. Nature 351:524–525.

Hall, E. R. 1981. The Mammals of North America. New York: Wiley.

Kolenosky, G. B., and R. O. Standfield. 1974. Morphological and ecological variation among gray wolves (Canis lupus) of Ontario, Canada. In M. W. Fox, ed., The Wild Canids: Their Systematics, Behavioral Ecology, and Evolution, 62–72. New York: Van Nostrand Reinhold.

Koler-Matznick, J., I. L. Brisbin Jr., M. Feinstein, and S. Bulmer. 2003. An updated description of the New Guinea singing dog (Canis hallstromi, Throughton 1957). Journal of Zoology 261:109–118.

Langguth, A. 1974. Ecology and evolution in the South American canids. In M. W. Fox, ed., The Wild Canids: Their Systematics, Behavioral Ecology, and Evolution, 192–206. New York: Van Nostrand Reinhold.

Larivière, S. 2002. Vulpes zerda. Mammalian Species 714:1–5.

Larivière, S., and M. Pasitschniak-Arts. 1996. Vulpes vulpes. Mammalian Species 537:1–11.

Larivière, S., and P. J. Seddon. 2001. Vulpes rueppelli. Mammalian Species 678:1–5.

Leonard, J. A., C. Vilà, and R. K. Wayne. 2005. Legacy lost: Genetic variability and population size of extirpated US grey wolves (Canis lupus). Molecular Ecology 14:9–17.

Macdonald, D. W., S. Creel, and M. G. L. Mills. 2004. Canid society. In D. W. Macdonald and C. Sillero-Zubiri, eds., The Biology and Conservation of Wild Canids, 85–106. Oxford: Oxford University Press.

Macdonald, D. W., and C. Sillero-Zubiri, eds. 2004. The Biology and Conservation of Wild Canids. Oxford: Oxford University Press.

MacIntosh, N. W. G. 1974. The origin of the dingo: An enigma. In M. W. Fox, ed., The Wild Canids: Their Systematics, Behavioral Ecology, and Evolution, 87–88. New York: Van Nostrand Reinhold.

Martensz, P. N. 1971. Observations on the food of the fox, Vulpes vulpes (L.), in an arid environment. Wildlife Research 16:73–75.

McGrew, J. C. 1979. Vulpes macrotis. Mammalian Species 123:1–6.

Mech, L. D. 1974. Canis lupus. Mammalian Species 37:1–6.

Mendelssohn, H., Y. Tom-Tov, G. Ilany, and D. Meninger. 1987. On the occurrence of Blanford’s fox, Vulpes cana Blanford, 1877, in Israel and Sinai. Mammalia 51:459–462.

Mivart, St. G. 1890. A Monograph of the Canidae: Dogs, Jackals, Wolves, and Foxes. London: Porter and Dulau.

Moore, C. M., and P. W. Collins. 1995. Urocyon littoralis. Mammalian Species 489:1–7.

Novaro, A. J. 1997. Pseudalopex culpaeus. Mammalian Species 558:1–8.

Nowak, R. M. 1992. The red wolf is not a hybrid. Conservation Biology 6:593–595.

Nowak, R. M. 2002. The original status of wolves in eastern North America. Southeastern Naturalist 1:95–130.

O’Brien, S. J., and E. Mayr. 1991. Bureaucratic mischief: Recognizing endangered species and subspecies. Science 251:1187–1188.

Osgood, W. H. 1934. The genera and subgenera of South American canids. Journal of Mammalogy 15:45–50.

Paradiso, J. L., and R. M. Nowak. 1971. A report on the taxonomic status and distribution of the red wolf. Fish and Wildlife Service Special Scientific Report 145:1–36.

Paradiso, J. L., and R. M. Nowak. 1972. Canis rufus. Mammalian Species 22:1–4.

Peres, C. A. 1991. Observations on hunting by small-eared (Atelocynus microtis) and bush dogs (Speothos venaticus) in central–western Amazonia. Mammalia 55:635–639.

Rook, L., and M. L. A. Puccetti. 1996. Remarks on the skull morphology of the endangered Ethiopian jackal, Canis simensis Rüppel 1838. Scienze Fisiche e Naturali 7:277–302.

Rosenzweig, M. L. 1966. Community structure in sympatric Carnivora. Journal of Mammalogy 47:602–612.

Servin, J. I. 1991. Algunos aspectos de la conducta social del lobo mexicano (Canis lupus baileyi) en cautiverio. Acta Zoologica Mexicana, n.s., 45:1–43.

Servin, J. I., and C. Huxley. 1991. La dieta del coyote en un bosque de encino-pino de la Sierra Madre Occidental de Durango, Mexico. Acta Zoologica Mexicana, n.s., 44:1–26.

Servin, J. I., J. R. Rau, and M. Delibes. 1987. Use of radio tracking to improve the estimation by track counts of the relative abundance of red fox. Acta Theriologica 32:489–492.

Servin, J. I., J. R. Rau, and M. Delibes. 1991. Activity pattern of the red fox Vulpes vulpes in Donana, SW Spain. Acta Theriologica 36:369–373.

Sillero-Zubiri, C., and D. Gottelli. 1994. Canis simensis. Mammalian Species 485:1–6.

Stains, H. J. 1975. Calcanea of members of the Canidae. Bulletin of the Southern California Academy of Sciences 74:143–155.

Strahl, S. D., J. L. Silva, and I. R. Goldstein. 1992. The bush dog (Speothos venaticus) in Venezuela. Mammalia 56:9–13.

Walton, L. R., and D. O. Joly. 2003. Canis mesomelas. Mammalian Species 715:1–9.

Ward, O. G., and D. H. Wurster-Hill. 1990. Nyctereutes procyonoides. Mammalian Species 358:1–5.

Wayne, R. K. 1986. Cranial morphology of domestic and wild canids: The influence of development on morphological change. Evolution 40:243–261.

Wayne, R. K., S. B. George, D. Gilbert, P. W. Collins, S. D. Kovach, D. J. Girman, and N. Lehman. 1991. A morphologic and genetic study of the island fox, Urocyon littoralis. Evolution 45:1849–1868.

Wilson, P. J., S. Grewal, I. D. Lawford, J. N. M. Heal, A. G. Granacki, D. Pennock, J. B. Theberge, M. T. Theberge, D. R. Voigt, W. Waddell, R. E. Chambers, P. C. Paquet, G. Goulet, D. Cluff, and B. N. White. 2000. DNA profiles of the eastern Canadian wolf and the red wolf provide evidence for a common evolutionary history independent of the gray wolf. Canadian Journal of Zoology 78:2156–2166.

Wozencraft, W. C. 1993. Order Carnivora. In D. E. Wilson and D. M. Reeder, eds., Mammal Species of the World: A Taxonomic and Geographic Reference, 2d ed., 279–348. Washington, D.C.: Smithsonian Institution Press.

Yahnke, C. J., W. E. Johnson, E. Geffen, D. Smith, F. Hertel, M. S. Roy, C. F. Bonacic, T. K. Fuller, B. Van Valkenburgh, and R. K. Wayne. 1996. Darwin’s fox: A distinct endangered species in a vanishing habitat. Conservation Biology 10:366–375.

Yeager, L. E. 1938. Tree-climbing by a gray fox. Journal of Mammalogy 19:376.

Zimmerman, R. S. 1938. A coyote’s speed and endurance. Journal of Mammalogy 19:400.

MOLECULAR BIOLOGY

Gottelli, D., C. Sillero-Zubiri, G. D. Applebaum, M. S. Roy, D. J. Girman, J. Garcia-Moreno, E. A. Ostrander, and R. K. Wayne. 1994. Molecular genetics of the most endangered canid: The Ethiopian wolf, Canis simensis. Molecular Ecology 3:277–290.

Lan, H., and L. Shi. 1996. The mitochondrial DNA evolution of four species of Canidae. Acta Zoologica Sinica 42:87–95.

Leonard, J. A., C. Vilà, K. Fox-Dobbs, P. L. Koch, R. K. Wayne, and B. Van Valkenburgh. 2007. Megafaunal extinctions and the disappearance of a specialized wolf Ecomorph. Current Biology 17:1146–1150.

Roy, M. S., E. Geffen, D. Smith, E. A. Ostrander, and R. K. Wayne. 1994. Patterns of differentiation and hybridization in North American wolflike canids, revealed by analysis of microsatellite loci. Molecular Biology and Evolution 11:553–570.

Wayne, R. K. 1993. Molecular evolution of the dog family. Trends in Genetics 9:218–224.

Wayne, R. K., E. Geffen, and C. Vilà. 2004. Population genetics: Population and conservation genetics of canids. In D. W. Macdonald and C. Sillero-Zubiri, eds., The Biology and Conservation of Wild Canids, 55–84. Oxford: Oxford University Press.

Wayne, R. K., and S. M. Jenks. 1991. Mitochondrial DNA analysis implying extensive hybridization of the endangered red wolf Canis rufus. Nature 351:565–568.

Wayne, R. K., W. G. Nash, and S. J. O’Brien. 1987a. Chromosomal evolution of the Canidae. I. Species with high diploid numbers. Cytogenetics and Cell Genetics 44:123–133.

Wayne, R. K., W. G. Nash, and S. J. O’Brien. 1987b. Chromosomal evolution of the Canidae. II. Divergence from the primitive carnivore karyotype. Cytogenetics and Cell Genetics 44:134–141.

Wayne, R. K., B. Van Valkenburgh, P. W. Kat, T. K. Fuller, W. E. Johnson, and S. J. O’Brien. 1989. Genetic and morphological divergence among sympatric canids. Journal of Heredity 80:447–454.

PALEOCLIMATE AND PALEOECOLOGY

Berggren, W. A., and D. R. Prothero. 1992. Eocene–Oligocene climatic and biotic evolution. In D. R. Prothero and W. A. Berggren, eds., Eocene–Oligocene Climatic and Biotic Evolution, 1–28. Princeton, N.J.: Princeton University Press.

Cavelier, C., J.-J. Chateauneuf, C. Pomerol, D. Rabussier, M. Renard, and C. V. Grazzini. 1981. The geological events at the Eocene/Oligocene boundary. Palaeogeography, Palaeoclimatology, Palaeoecology 36:223–248.

Cerling, T. E. 1992. Development of grasslands and savannas in East Africa during the Neogene. Palaeogeography, Palaeoclimatology, Palaeoecology 97:241–247.

Cerling, T. E., J. Quade, Y. Wang, and J. R. Bowman. 1989. Carbon isotopes in soils and paleosols as ecology and paleoecology indicators. Nature 341:138–139.

Crusafont-Pairó, M., and J. Truyols-Santonja. 1956. A biometric study of the evolution of fissiped carnivores. Evolution 10:314–332.

DeConto, R. M., and D. Pollard. 2003. Rapid Cenozoic glaciation of Antarctica induced by declining atmospheric CO₂. Nature 421:245–249.

Fortelius, M., J. Eronen, J. Jernvall, L. Liu, D. Pushkina, J. Rinne, A. Tesakov, I. A. Vislobokova, Z. Zhang, and L. Zhou. 2002. Fossil mammals resolve regional patterns of Eurasian climate change over 20 million years. Evolutionary Ecology Research 4:1005–1016.

Fortelius, M., and N. Solounias. 2000. Functional characterization of ungulate molars using the abrasion-attrition wear gradient: A new method for reconstructing paleodiets. American Museum Novitates 3301:1–36.

Hu, Y., J. Meng, Y. Wang, and C. Li. 2005. Large Mesozoic mammals fed on young dinosaurs. Nature 433:149–152.

Hunt, R. M., Jr. 2004. Global climate and the evolution of large mammalian carnivores during the later Cenozoic in North America. Bulletin of the American Museum of Natural History 285:139–156. [Special issue: G. C. Gould and S. K. Bell, eds., Tributes to Malcolm C. McKenna: His Students, His Legacy]

Janis, C. M., J. Damuth, and J. M. Theodor. 2002. The origins and evolution of the North American grassland biome: The story from the hoofed mammals. Palaeogeography, Palaeoclimatology, Palaeoecology 177:183–198.

Kennett, J. P. 1977. Cenozoic evolution of Antarctic glaciation, the circum-Antarctic oceans and their impact on global paleoceanography. Journal of Geophysical Research 82:3843–3860.

Leopold, E. B., G. Liu, and S. Clay-Poole. 1992. Low-biomass vegetation in the Oligocene? In D. R. Prothero and W. A. Berggren, eds., Eocene–Oligocene Climatic and Biotic Evolution, 399–420. Princeton, N.J.: Princeton University Press.

MacFadden, B. J. 2000. Cenozoic mammalian herbivores from the Americas: Reconstructing ancient diets and terrestrial communities. Annual Review of Ecology and Systematics 31:33–59.

MacFadden, B. J., and T. E. Cerling. 1994. Fossil horses, carbon isotopes, and global change. Trends in Ecology and Evolution 9:481–486.

MacFadden, B. J., and T. E. Cerling. 1996. Mammalian herbivore communities, ancient feeding ecology, and carbon isotopes: A 10-million-year sequence from the Neogene of Florida. Journal of Vertebrate Paleontology 16:103–115.

Palmqvist, P., A. Arribas, and B. Martínez-Navarro. 1999. Ecomorphological study of large canids from the lower Pleistocene of southeastern Spain. Lethaia 32:75–88.

Palmqvist, P., B. Martínez-Navarro, and A. Arribas. 1996. Prey selection by terrestrial carnivores in a lower Pleistocene paleocommunity. Paleobiology 22:514–534.

Pearson, P. M., P. W. Ditchfield, J. Singano, K. G. Harcourt-Brown, C. J. Nicholas, R. K. Olsson, N. J. Shackleton, and M. A. Hall. 2001. Warm tropical sea surface temperatures in the late Cretaceous and Eocene epochs. Nature 413:481–487.

Prothero, D. R., and T. H. Heaton. 1996. Faunal stability during the early Oligocene climatic crash. Palaeogeography, Palaeoclimatology, Palaeoecology 127:257–283.

Quade, J., and T. E. Cerling. 1995. Expansion of C4 grasses in the late Miocene of northern Pakistan: Evidence from stable isotopes in paleosols. Palaeogeography, Palaeoclimatology, Palaeoecology 115:91–116.

Ramstein, G., F. Fluteau, J. Bese, and S. Joussaume. 1997. Effect of orogeny, plate motion, and land-sea distribution on Eurasian climate change over the past 30 million years. Nature 386:788–795.

Ridgway, K. D., A. R. Sweet, and A. R. Cameron. 1995. Climatically induced floristic changes across the Eocene–Oligocene transition in the northern high latitudes, Yukon Territory, Canada. Bulletin of the Geological Society of America 107:676–696.

Solounias, N., and G. Semprebon. 2002. Advances in the reconstruction of ungulate ecomorphology with application to early fossil equids. American Museum Novitates 3366:1–49.

Strömberg, C. A. E. 2002. The origin and spread of grass-dominated ecosystems in the late Tertiary of North America: Preliminary results concerning the evolution of hypsodonty. Palaeogeography, Palaeoclimatology, Palaeoecology 177:59–75.

Van Valkenburgh, B. 1994. Extinction and replacement among predatory mammals in the North American late Eocene–Oligocene: Tracking a guild over twelve million years. Historical Biology 8:1–22.

Van Valkenburgh, B. 1999. Major patterns in the history of carnivorous mammals. Annual Review of Earth and Planetary Science 27:463–493.

Van Valkenburgh, B. 2001. The dog-eat-dog world of carnivores: A review of past and present carnivore community dynamics. In C. Stanford and H. T. Bunn, eds., Meat-Eating and Human Evolution, 101–121. Oxford: Oxford University Press.

Van Valkenburgh, B., and F. Hertel. 1993. Tough times at La Brea: Tooth breakage in large carnivores of the late Pleistocene. Science 261:456–459.

Van Valkenburgh, B., and T. Sacco. 2002. Sexual dimorphism, social behavior, and intrasexual competition in large Pleistocene carnivorans. Journal of Vertebrate Paleontology 22:164–169.

Van Valkenburgh, B., X. Wang, and J. Damuth. 2004. Cope’s rule, hypercarnivory, and extinction in North American canids. Science 306:101–104.

Wang, Y., and T. E. Cerling. 1994. A model of fossil tooth and bone diagenesis: Implications for paleodiet reconstruction from stable isotopes. Palaeogeography, Palaeoclimatology, Palaeoecology 107:281–289.

Wang, Y., T. E. Cerling, and B. J. MacFadden. 1994. Fossil horses and carbon isotopes: New evidence for Cenozoic dietary, habitat, and ecosystem changes in North America. Palaeogeography, Palaeoclimatology, Palaeoecology 107:269–279.

Webb, S. D. 1977. A history of savanna vertebrates in the New World. Part I: North America. Annual Review of Ecology and Systematics 8:355–380.

Wing, S. L., G. J. Harrington, F. A. Smith, J. I. Bloch, D. M. Boyer, and K. H. Freeman. 2005. Transient floral change and rapid global warming at the Paleocene–Eocene boundary. Science 310:993–996.

Zachos, J. C., M. Pagani, L. Sloan, and E. Thomas. 2001. Trends, rhythms, and aberrations in global climate 65 Ma to present. Science 292:686–693.

Zachos, J. C., L. D. Stott, and K. C. Lohmann. 1994. Evolution of early Cenozoic marine temperatures. Paleoceanography 9:353–387.

PHYLOGENY, SYSTEMATICS, AND TAXONOMY

Bardeleben, C., R. L. Moore, and R. K. Wayne. 2005. A molecular phylogeny of the Canidae based on six nuclear loci. Molecular Phylogenetics and Evolution 37:815.

Clutton-Brock, J., G. B. Corbet, and M. Hills. 1976. A review of the family Canidae, with a classification by numerical methods. Bulletin of the British Museum (Natural History), Zoology 29:119–199.

Dahr, E. 1949. On the systematic position of Phlaocyon leucosteus Matthew and some related forms. Arkiv för Zoologi 41A:1–15.

Flynn, J. J., N. A. Neff, and R. H. Tedford. 1988. Phylogeny of the Carnivora. In M. J. Benton, ed., The Phylogeny and Classification of the Tetrapods, vol. 2, Mammals, 73–116. Oxford: Clarendon Press.

Geffen, E., A. Mercure, D. J. Girman, D. W. Macdonald, and R. K. Wayne. 1992. Phylogenetic relationships of the fox-like canids: Mitochondrial DNA restriction fragment, site, and cytochrome b sequence analyses. Journal of Zoology 228:27–39.

Grewal, S., P. J. Wilson, T. K. Kung, K. Shami, M. T. Theberge, J. B. Theberge, and B. N. White. 2004. A genetic assessment of the eastern wolf (Canis lycaon) in Algonquin Provincial Park. Journal of Mammalogy 85:625–632.

Hough, J. R. 1944. The auditory region in some Miocene carnivores. Journal of Paleontology 22:573–600.

Hough, J. R. 1948. The auditory region in some members of the Procyonidae, Canidae, and Ursidae. Bulletin of the American Museum of Natural History 92:73–118.

Hunt, R. M., Jr. 1974. The auditory bulla in Carnivora: An anatomical basis for reappraisal of carnivore evolution. Journal of Morphology 143:21–76.

Huxley, T. H. 1880. On the cranial and dental characters of the Canidae. Proceedings of the Zoological Society of London 16:238–288.

Lawrence, B., and W. H. Bossert. 1974. Relationships of North American Canis shown by a multiple character analysis of selected populations. In M. W. Fox, ed., The Wild Canids: Their Systematics, Behavioral Ecology, and Evolution, 73–86. New York: Van Nostrand Reinhold.

Ledje, C., and U. Arnason. 1996a. Phylogenetic analyses of complete cytochrome b genes of the order Carnivora with particular emphasis on the Caniformia. Journal of Molecular Evolution 42:135–144.

Ledje, C., and U. Arnason. 1996b. Phylogenetic relationships within caniform carnivores based on analyses of the mitochondrial 12S rRNA gene. Journal of Molecular Evolution 43:641–649.

Lyras, G. A., and A. A. E. Van der Geer. 2003. External brain anatomy in relation to the phylogeny of Canidae (Carnivora: Canidae). Zoological Journal of the Linnean Society 138:505–522.

Martin, L. D. 1989. Fossil history of the terrestrial Carnivora. In J. L. Gittleman, ed., Carnivore Behavior, Ecology, and Evolution, 1:536–568. Ithaca, N.Y.: Cornell University Press.

Matthew, W. D. 1930. The phylogeny of dogs. Journal of Mammalogy 11:117–138.

Mayr, E. 1940. Speciation phenomena in birds. American Naturalist 74:249–278.

Mayr, E. 1969. Principles of Systematic Zoology. New York: McGraw-Hill.

McKenna, M. C., and S. K. Bell. 1997. Classification of Mammals Above the Species Level. New York: Columbia University Press.

Nowak, R. M. 1979. North American Quaternary Canis. Monograph of the Museum of Natural History, University of Kansas 6:1–154.

Phillips, M., and V. G. Henry. 1992. Comments on red wolf taxonomy. Conservation Biology 6:596–599.

Segall, W. 1943. The auditory region of the arctoid carnivores. Field Museum of Natural History, Zoological Series 29:33–59.

Simpson, G. G. 1945. The principles of classification and a classification of mammals. Bulletin of the American Museum of Natural History 8:1–350.

Stains, H. J. 1974. Distribution and taxonomy of the Canidae. In M. W. Fox, ed., The Wild Canids: Their Systematics, Behavioral Ecology, and Evolution, 3–26. New York: Van Nostrand Reinhold.

Tedford, R. H. 1976. Relationship of pinnipeds to other carnivores (Mammalia). Systematic Zoology 25:363–374.

Tedford, R. H., B. E. Taylor, and X. Wang. 1995. Phylogeny of the Caninae (Carnivora: Canidae): The living taxa. American Museum Novitates 3146:1–37.

Turner, H. N., Jr. 1848. Observations relating to some of the foramina at the base of the skull in Mammalia, and on the classification of the order Carnivora. Proceedings of the Zoological Society of London 16:63–88.

Van Gelder, R. G. 1978. A review of canid classification. American Museum Novitates 2646:1–10.

Wang, X., and R. H. Tedford. 1994. Basicranial anatomy and phylogeny of primitive canids and closely related miacids (Carnivora: Mammalia). American Museum Novitates 3092:1–34.

Wang, X., R. H. Tedford, and B. E. Taylor. 1999. Phylogenetic systematics of the Borophaginae (Carnivora: Canidae). Bulletin of the American Museum of Natural History 243:1–391.

Wayne, R. K. 1993. Molecular evolution of the dog family. Trends in Genetics 9:218–224.

Wayne, R. K., R. E. Benveniste, D. N. Janczewski, and S. J. O’Brien. 1989. Molecular and biochemical evolution of the Carnivora. In J. L. Gittleman, ed., Carnivore Behavior, Ecology, and Evolution, 1:465–494. Ithaca, N.Y.: Cornell University Press.

Wayne, R. K., E. Geffen, D. J. Girman, K.-P. Koepfli, L. M. Lau, and C. R. Marshall. 1997. Molecular systematics of the Canidae. Systematic Zoology 46:622–653.

Wayne, R. K., and S. J. O’Brien. 1987. Allozyme divergence within the Canidae. Systematic Zoology 36:339–355.

Wozencraft, W. C. 1989. The phylogeny of the recent Carnivora. In J. L. Gittleman, ed., Carnivore Behavior, Ecology, and Evolution, 1:495–535. Ithaca, N.Y.: Cornell University Press.

Zrzavý, J., and V. Ricánková. 2004. Phylogeny of recent Canidae (Mammalia, Carnivora): Relative reliability and utility of morphological and molecular datasets. Zoologica Scripta 33:311–333.

Zunino, G. E., O. B. Vaccaro, M. Canevari, and A. L. Gardner. 1995. Taxonomy of the genus Lycalopex (Carnivora: Canidae) in Argentina. Proceedings of the Biological Society of Washington 108:729–747.

SEXUAL DIMORPHISM

Gingerich, P. D., and D. A. Winkler. 1979. Patterns of variation and correlation in the dentition of the red fox, Vulpes vuples. Journal of Mammalogy 60:691–704.

Gittleman, J. L., and B. Van Valkenburgh. 1997. Sexual dimorphism in the canines and skulls of carnivores: Effects of size, phylogeny, and behavioural ecology. Journal of Zoology 242:97–117.

Jolicoeur, P. 1974. Sexual dimorphism and geographical distance as factors of skull variation in the wolf Canis lupus L. In M. W. Fox, ed., The Wild Canids: Their Systematics, Behavioral Ecology, and Evolution, 54–61. New York: Van Nostrand Reinhold.

Prestrud, P., and K. Nilssen. 1995. Growth, size, and sexual dimorphism in arctic foxes. Journal of Mammalogy 76:522–530.

Regodon, S., A. Franco, J. M. Garin, A. Robina, and Y. Lignereux. 1991. Computerized tomographic determination of the cranial volume of the dog applied to racial and sexual differentiation. Acta Anatomica 142:347–350.

ZOOGEOGRAPHY

Berta, A. 1987. Origin, diversification, and zoogeography of the South American Canidae. In B. D. Patterson and R. M. Timm, eds., Studies in Neotropical Mammalogy: Essays in Honor of Philip Hershkovitz, 455–471. Fieldiana Zoology, n.s., 39. Chicago: Field Museum of Natural History.

Brunet, M., F. Guy, D. Pilbeam, D. E. Lieberman, A. Likius, H. T. Mackaye, M. S. Ponce de Leon, C. P. E. Zollikofer, and P. Vignaud. 2005. New material of the earliest hominid from the upper Miocene of Chad. Nature 434:752–755.

Hunt, R. M., Jr. 1996. Biogeography of the order Carnivora. In J. L. Gittleman, ed., Carnivore Behavior, Ecology, and Evolution, 2:485–541. Ithaca, N.Y.: Cornell University Press.

Johnson, W. E., T. K. Fuller, and W. L. Franklin. 1989. Sympatry in canids: A review and assessment. In J. L. Gittleman, ed., Carnivore Behavior, Ecology, and Evolution, 1:189–217. Ithaca, N.Y.: Cornell University Press.

Qiu, Z. 2003. Dispersals of Neogene carnivorans between Asia and North America. Bulletin of the American Museum of Natural History 279:18–31.

Stains, H. J. 1974. Distribution and taxonomy of the Canidae. In M. W. Fox, ed., The Wild Canids: Their Systematics, Behavioral Ecology, and Evolution, 3–26. New York: Van Nostrand Reinhold.

Vilà, C., I. R. Amorim, J. A. Leonard, D. Posada, J. Castroviejo, F. Petrucci-Fonseca, K. A. Crandall, H. Ellegren, and R. K. Wayne. 1999. Mitochondrial DNA phylogeography and population history of the grey wolf Canis lupus. Molecular Ecology 8:2089–2103.