Abstract
Analyses of anatomical and DNA sequence data run on a parallel supercomputer that include fossil taxa support the inclusion of tenrecs and golden moles in the Afrotheria, an endemic African clade of placental mammals. According to weighting schemes of morphological and molecular data that maximize congruence, extinct members of the afrotherian crown group include embrithopods, Plesiorycteropus, desmostylians, and the “condylarths” Hyopsodus, Meniscotherium, and possibly Phenacodus. By influencing the optimization of anatomical characters, molecular data have a large influence on the relationships of several extinct taxa. The inclusion of fossils and morphological data increases support for an elephant-sea cow clade within Paenungulata and identifies ancient, northern elements of a clade whose living members in contrast suggest an historically Gondwanan distribution. In addition, maximally congruent topologies support the position of Afrotheria as well-nested, not basal, within Placentalia. This pattern does not accord with the recent hypothesis that the divergence of placental mammals co-occurred with the tectonic separation of Africa and South America.
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Literature Cited
Adkins, R. M., Walton, A. H., and Honeycutt, R. L. (2003). Higher-level systematics of rodents and divergence time estimates based on two congruent nuclear genes. Mol. Phylogenet. Evol. 26: 409–420.
Agur, A. M. R., and Lee, M. J. (1991). Grant's Atlas of Anatomy, 9th edn. Williams and Wilkins, Baltimore.
Amrine, H. M., and Springer, M. S. (1999). Maximum likelihood analysis of the tethythere hypothesis. J. Mammal. Evol. 6: 161–176.
Andrews, C. W. (1906). A Descriptive Catalogue of the Tertiary Vertebrata of the Fayum, Egypt, British Museum of Natural History, London.
Arnason, U., Addegoke, J. A., Bodin, K., Born, E. W., Esa, Y. B., Gullberg, A., Nilsson, M., Short, R., Xu, X., and Janke, A. (2002). Mammalian mitogenomic relationships and the root of the eutherian tree. Proc. Natl. Acad. Sci. U.S.A. 99: 8151–8156.
Asher, R. J. (1999). A morphological basis for assessing the phylogeny of the “Tenrecoidea” (Mammalia, Lipotyphla). Cladistics 15: 231–252.
Asher, R. J. (2000). Phylogenetic History of Tenrecs and Other Insectivoran Mammals. Ph.D. Dissertation, State University of New York, Stony Brook.
Asher, R. J. (2001). Cranial anatomy in tenrecid insectivorans: Character evolution across competing phylogenies. Amer. Mus. Novit. 3352: 1–54.
Asher, R. J. (2003). Phylogenetics of the Tenrecidae (Mammalia): A response to Douady et al., 2002. Mol. Phylogenet. Evol. 26: 328–330.
Barnes, L. G. (1990). The fossil record and evolutionary relationships of the genus Tursiops. In: The Bottlenosed Dolphin, S. Leatherwood and R. R. Reeves, eds., pp. 3–26, Academic Press, New York.
Beard, K. C., and MacPhee, R. D. E. (1994). Cranial anatomy of Shoshonius and the antiquity of the Anthropoidea. In: Anthropoid Origins, J. G. Fleagle and R. F. Kay, eds., pp. 55–97, Plenum, New York.
Butler, P. M. (1956). The skull of Ictops and the classification of the Insectivora. Proc. Zool. Soc. Lond. 126: 453–481.
Butler, P. M. (1988). Phylogeny of the insectivores. In: Phylogeny and Classification of the Tetrapods, Vol. 2, M. J. Benton, ed., pp. 117–141, Clarendon Press, Oxford.
Carter, A. M. (2001). Evolution of the placenta and fetal membranes seen in the light of molecular phylogenetics. Placenta 22: 800–807.
Cartmill, M. (1978). The orbital mosaic in prosimians and the use of variable traits in systematics. Folia Primatol. 30(2): 89–114.
Cartmill, M. (1980). Morphology, function, and evolution of the anthropoid postorbital septum. In: Evolutionary Biology of the New World Monkeys and Continental Drift. R. L. Ciochon and A. B. Chiarelli, eds., pp. 243–274, Plenum, New York.
Cartmill, M., and MacPhee, R. D. E. (1980). Tupaiid affinities: The evidence of the carotid arteries and cranial skeleton. In: Comparative Biology and Evolutionary Relationships of Tree Shrews, P. Luckett, ed., pp. 95–132, Plenum, New York.
Cifelli, R. (1982). The petrosal structure of Hyopsodus with respect to that of some other ungulates, and its phylogenetic implications. J. Paleontol. 56: 796–805.
Clark, J. M. (1991). A new early Miocene species of Paleoparadoxia (Mammalia: Desmostylia) from California. J. Vertebr. Paleontol. 11: 490–508.
Colless, D. H. (1980). Congruence between morphometric and allozyme data for Menidia species: A reappraisal. Syst. Zool. 29: 288–299.
Court, N. (1990). Periotic anatomy of Arsinoitherium (Mammalia, Embrithopoda) and its phylogenetic implications. J. Vertebr. Paleontol. 10: 170–182.
Court, N. (1992). The skull of Arsinoitherium (Mammalia, Embrithopoda) and the higher order phylogenetic relationships of ungulate mammals. Palaeovertebrata (Montpelier) 22: 1–43.
Court, N. (1993). Morphology and functional anatomy of the postcranial skeleton in Arsinoitherium (Mammalia, Embrithopoda). Palaeontographica (Stuttgart) A 226: 125–169.
Court, N. (1994). The periotic of Moeritherium (Mammalia, Proboscidea): Homology or homoplasy in the ear region of Tethytheria McKenna, 1975. Zool. J. Linn. Soc. 112: 13–28.
Court, N. (1995). A new species of Numidotherium (Mammalia, Proboscidea) from the Eocene of Libya and the early phylogeny of the Proboscidea. J. Vertebr. Paleontol. 15: 650–671.
Dathe, F. (1982). Megaptera hubachi n. sp., ein fossiler Bartenwahl aus marinen Sandsteinschichten des tieferen Pliozäns Chiles. Z. Geol. Wiss. 11: 813–848.
Domning, D., Ray, C. E., and McKenna, M. C. (1986). Two new OIigocene desmostylians and a discussion of tethytherian systematics. Smithson. Contrib. Paleobiol. 59: 1–56.
Douady, C. J., Catzeflis, F., Kao, D. J., Springer, M. S., and Stanhope, M. J. (2002a). Molecular evidence for the monophyly of tenrecidae (Mammalia) and the timing of the colonization of Madagascar by Malagasy tenrecs. Mol. Phylogenet. Evol. 22: 357–363.
Douady, C. J., Chatelier, P. I., Madsen, O., de Jong, W. W., Catzeflis, F., Springer, M. S., and Stanhope, M. J. (2002b). Molecular phylogenetic evidence confirming the Eulipotyphla concept and in support of hedgehogs as the sister group to shrews. Mol. Phylogenet. Evol. 25: 200–209.
Douady, C. J., Catzeflis, F., Springer, M. S., and Stanhope, M. J. (2003). Phylogenetics of the Tenrecidae (Mammalia): A reply to Asher. Mol. Phylogenet. Evol. 26: 331–332.
Downton, M., and Austin, A. D. (2002). Increased congruence does not necessarily indicate increased phylogenetic accuracy. Syst. Biol. 51: 19–31.
Emerson, G. L., Kilpatrick, C. W., McNiff, B. E., Ottenwalder, J., and Allard, M. W. (1999). Phylogenetic relationships of the order Insectivora based on complete 12s rRNA sequences from mitochondria. Cladistics 15: 221–230.
Felsenstein, J. (1978). Cases in which parsimony or compatibility methods will be positively misleading. Syst. Zool. 27: 401–410.
Felsenstein, J. (1988). Phylogenies from molecular sequences: Inference and reliability. Annu. Rev. Genet. 22: 521–565.
Fischer, M. S. (1986). Die Stellung der Schliefer (Hyracoidea) im phylogenetischen System der Eutheria. Cour. Forsch. Inst. Senckenberg. 84: 1–132.
Fischer, M. S. (1998). Die Lokomotion von Procavia capensis (Mammalia: Hyracoidea): zur Evolution des Bewegungssystems bei Saugetieren. Abhandlungen des Naturwissenschaftlichen Vereins in Hamburg. 33: 1–188.
Fischer, M., and P. Tassy. (1993). The interrelation between Proboscidea, Sirenia, Hyracoidea, and Mesaxonia: The morphological evidence. In: Mammal Phylogeny, Vol. 2: Placentals, F. S. Szalay, M. J. Novacek, and M. C. McKenna, eds., pp. 217–234, Springer-Verlag, New York.
Fleagle, J. G. (1999). Primate Adaptation and Evolution, 2nd edn. Academic Press, San Diego.
Frost, D. R., Rodrigues, M. T., Grant, T., and Titus, T. (2001). Phylogenetics of the lizard genus Tropidurus (Squamata: Tropiduridae). Mol. Phylogenet. Evol. 21: 352–371.
Frost, D. R., Wozencraft, C., and Hoffman, R. S. (1991). Phylogenetic relationships of hedgehogs and gymnures (Mammalia: Insectivora: Erinaceidae). Smithson. Contrib. Zool. 518: 1–69.
Gatesy, J., DeSalle, R., and Wheeler, W. (1993). Alignment ambiguous nucleotide sites and the exclusion of systematic data. Mol. Phylogenet. Evol. 2: 152–157.
Gatesy, J., Mathee, C., DeSalle, R., Hayashi, C. (2002). Resolution of a Supertree/Supermatrix Paradox. Syst. Biol. 51: 652–664.
Gaudin, T. J., Wible, J. R., Hopson, J. A., and Turnbull, W. D. (1996). Reexamination of the morphological evidence for the cohort Epitheria (Mammlia, Eutheria) J. Mammal. Evol. 3: 31–79.
Gazin, C. L. (1956). A study of the Eocene condylarthran mammal Hyopsodus. Smithson. Misc. Collect. 131: 1–57.
Gazin, C. L. (1965). A study of the early Tertiary condylarthran mammal Meniscotherium. Smithson. Misc. Collect. 149: 1–98.
Gazin, C. L. (1968). A study of the Eocene condylarthran mammal Hyopsodus. Smithson. Misc. Collect. 153: 1–90.
Geisler, J. H. (2001). New morphological evidence for the phylogeny of Artiodactyla, Cetacea, and Mesonychidae. Amer. Mus. Novit. 3344: 1–53.
Geisler, J. H., and Luo, Z. (1998). Relationships of Cetacea to terrestrial ungulates and the evolution of ranial vasculature in Cete. In: The Emergence of Whales, J. G. M. Thewissen, ed., pp. 163–212, Plenum, New York.
Gheerbrandt, E., Sudre, J., Iarochene, M., and Moumni, A. (2001). First ascertained African “condylarth” mammals from the earliest Ypresian of the Ouled Abdoun Basin, Morocco. J. Vertebr. Paleontol. 21: 107–118.
Giere, P. (2002). Grundplan rekonstruktion und Ontogenese der Orbitalregion der “Eulipotyphla” (Mammalia). PhD Thesis, Humboldt Univ. Wissenschaft und Technik Verlag, Berlin.
Giribet, G., Edgecomb, G. D., and Wheeler, W.C. (2001). Arthropod phylogeny based on eight molecular loci and morphology. Nature 413: 157–161.
Goloboff, P. A. (1999). NONA, Version 2.0. Fundación e Instituto Miguel Lillo, Tucumán, Argentina. Download at www.cladistics.org.
Gregory, W. K. (1910). The orders of mammals. Bull. Amer. Mus. Nat. Hist. 27: 1–524.
Grzimek, B. (1975). Grzimek's Animal Life Encyclopedia, I–IV, Van Nostrand Rheinhold, New York.
Hermanson, J. W., and MacFadden, B. J. (1996). Evolutionary and functional morphology of the knee in fossil and extant horses (Equidae). J. Vertebr. Paleontol. 16: 349–357.
Horovitz, I. (1999). A phylogenetic study of living and fossil platyrrhines. Amer. Mus. Novit. 3269: 1–40.
Horovitz, I. (2000). The tarsus of Ukhaatherium nessovi (Eutheria, Mammalia) from the Late Cretaceous of Mongolia: An appraisal of the evolution of the ankle in basal therians. J. Vertebr. Paleontol. 20: 547–560.
Ijiri, S., and Kamei, T. (1961). On the skulls of Desmostylus mirabilis Nagao from South Sakhalin and of Palaeoparadoxia tabatai Tokunaga from Gifu prefecture, Japan [in Japanese]. Earth Sci. 53: 1–27.
Janies, D. A., and Wheeler, W.C. (2001). Efficiency of parallel direct optimization. Cladistics 17: S71-S82.
Johnson, P. A., and Fox, R. C. (1984). Paleocene and Late Cretaceous mammals from Saskatchewan, Canada. Palaeontographica Abt. A. 186: 163–222.
Kamei, T., and Okazaki, Y. (1975). Neogene desmostylid and proboscidean fossils from Japan. Atlas and Jpn. Fossils 34(199 and 201): 1–4.
Kielan-Jaworowska, Z. (1979). Evolution of the therian mammals in the Late Cretaceous of Asia. Part III. Postcranial skeleton in Zalambdalestidae. Palaeontol. Pol. 38: 3–41.
Kielan Jaworowska, Z. (1984). Evolution of the therian mammals in the Late Cretaceous of Asia. Part V. Skull structure in Zalambdalestidae Palaeontol. Pol. 46: 107–117.
Krause, D. W., Rodgers, R. R., Forster, C. A., Hartman, J. H., Buckley, G. A., and Sampson, S. D. (1999). Late Cretaceous vertebrate fauna of Madagascar: Implications for Gondwanan paleobiogeography. GSA Today 9: 1–7.
Lee, M. S. Y. (2001). Unalignable sequences and molecular evolution. Trends Ecol. Evol. 16: 681–685.
Lin, Y.-H., McLeanachan, P. A., Gore, A. R., Phillips, M. J., Ota, R., Hendy, M. D., and Penny, D. (2002a). Four new mitochondrial genomes and the increased stability of evolutionary trees of mammals from improved taxon sampling. Mol. Biol. Evol. 19: 2060–2070.
Lin, Y.-H., Waddell, P., and Penny, D. (2002b). Pika and vole mitochondrial genomes increase support for both rodent monophyly and Glires. Gene 294: 119–129.
Lillegraven, J. A. (1969). Latest Cretaceous mammals of upper part of Edmonton Formation of Alberta, Canada, and review of marsupial-placental dichotomy in mammalian evolution. Univ. Kansas Paleontol. Contrib. 50(vertebrata 12): 1–122.
Lillegraven, J. A. (1984). Why was there a marsupial-placental dichotomy? In: Mammals: Notes from a Short Course, Vol. 8: Studies in Geology, P. D. Gingerich and C. E. Badgley, eds., pp. 72–86, University of Tennessee, Department of Geological Science, Knoxville.
Lillegraven, J. A. (1985). Use of the term “trophoblast” for tissues in therian mammals. J. Morphol. 183: 293–299.
Luckett, W. P. (1977). Ontogeny of amniote fetal membrances and their application to phylogeny. In: Major Patterns in Vertebrate Evolution, M. K. Hecht, P. C. Goody, and B. M. Hecht, eds., pp. 439–516, Plenum, New York.
Luckett, W. P. (1985). Superordinal and intraordinal affinities of rodents: Developmental evidence from the dentition and placentation. In: Evolutionary Relationships Among Rodents, W. P. Luckett and J.-L. Hartenberger, eds., pp. 227–276, Plenum, New York.
Luckett, W. P. (1993). Uses and limitations of mammalian fetal membrances and placenta for phylogenetic reconstruction. J. Exp. Zool. 266: 514–527.
Luo, Z. (1998). Homology and transformation of cetacean ectotympanic structures. In: The Emergence of Whales, J. G. M. Thewissen, ed., pp. 269–301, Plenum, New York.
Luo, Z., and Gingerich, P. D. (1999). Terrestrial Mesonychia to aquatic Cetacea: Transformation of the basicranium and evolution of hearing in whales. Univ. Mich. Pap. Paleontol. 31: 1–98.
MacPhee, R. D. E. (1979.) Entotympanics, ontogeny, and primates. Folia Primatol. 31: 23–47.
MacPhee, R. D. E. (1994). Morphology, adaptations, and relationships of Plesiorycteropus, and a diagnosis of a new order of eutherian mammals. Bull. Amer. Mus. Nat. Hist. 220: 1–214.
MacPhee, R. D. E., and Novacek, M. J. (1993). Definition and relationships of Lipotyphla. In: Mammal Phylogeny, Vol. 2: Placentals, F. S. Szalay, M. J. Novacek, and M. C. McKenna, eds., pp. 13–31, Springer-Verlag, New York.
MacPhee, R. D. E., Novacek, M. J., and Storch, G. (1988). Basicranial morphology of Early Tertiary erinaceomorphs and the origin of primates. Amer. Mus. Novit. 2921: 1–42.
Madsen, O., Scally, M., Douady, C. J., Kao, D. J., DeBry, R. W., Adkins, R., Amrine, H. M., Stanhope, M. J., de Jong, W. W., and Springer, M. S. (2001). Parallel adaptive radiations in two major clades of placental mammals. Nature 409: 610–614.
Mahboubi, M., Ameur, J. Y, Crochet, Y., and Jaeger, J. J. (1986). El Kohol (Saharan Atlas, Algeria), a new Eocene mammal locality in northwestern Africa. Palaeontographica Abt. A 192: 15–49.
Malia, M. J., Adkins, R. M., and Allard, M. W. (2002). Molecular support for Afrotheria and the polyphyly of Lipotyphla based on analyses of the growth hormone receptor gene. Mol. Phylogenet. Evol. 24: 91–101.
Matsumoto, H. (1923). A contribution to the knowledge of Moeritherium. Bull. Amer. Mus. Nat. Hist. 48: 97–139.
McDowell, S. B., Jr. (1958). The Greater Antillean insectivores. Bull. Amer. Mus. Nat. Hist. 115: 113–214.
McKenna, M. C. (1963). New evidence against tupaioid affinities of the mammlian family Anagalidae. Amer. Mus. Novit. 2158: 1–16.
McKenna, M. C., and BellS. K. (1997). Classification of Mammals Above the Species Level, Columbia University Press, New York.
McKenna, M. C., and Manning, R. (1977). Affinities and palaeobiogeographic significance of the Mongolian Paleogene genus Phenacolophus. Geobios Mem. Spec. 1: 61–85.
Meng, J., Hu, Y., and Li, C. (2003). The osteology of Rhombomylus (Mammalia: Glires): Implications for the phylogeny and evolution of Glires. Bull. Amer. Mus. Nat. Hist. 275: 1–247.
Meng, J., and Wyss, A. (2001). The morphology of Tribosphenomys (Rodentiaformes, Mammalia): Phylogenetic implications for basal Glires.J. Mammal. Evol. 8: 1–72.
Millar, J. S. (1981). Pre-partum reproductive characteristics of eutherian mammals. Evolution. 35: 1149–1163.
Mossman, H. W. (1987). Vertebrate Fetal Membranes, Rutgers University Press, New Brunswick.
Mouchaty, S. K., Gullberg, A., Janke, A., and Arnason, U. (2000). Phylogenetic position of the tenrecs (Mammalia: Tenrecidae) of Madagascar based on analysis of the complete mitochondrial genome sequence of Echinops telfairi. Zoologica Scripta 29: 307–317.
Murphy, W. J., Elzirik, E., Johnson, W. E., Zhang, Y. P., Ryder, O. A., and O'Brien, S. J. (2001a). Molecular phylogenetics and the origin of placental mammals. Nature 409: 614–618.
Murphy, W. J., Eizirik, E., O'Brien, S. J., Madsen, O., Scally, M., Douady, C. J., Teeling, E., Ryder, O. A., Stanhope, M. J., deJong, W. W., and Springer, M. S. (2001b). Resolution of the early placental mammal radiation using Baysian phylogenetics. Science 294: 2348–2351.
Novacek, M. J. (1977). Aspects of the problem of variation, origin, and evolution of the eutherian auditory bulla. Mammal. Rev. 7: 131–149.
Novacek, M. J. (1986). The skull of leptictid insectivorans and the higher-level classification of eutherian mammals. Bull. Amer. Mus. Nat. Hist. 183: 1–111.
Novacek, M. J. (1989). Higher mammal phylogeny: The morphological-molecular synthesis. In: The Hierarchy of Life, B. Fernholm, K. Bremer, and H. Jörnvall, H., eds., pp. 421–435, Elsevier, Amsterdam.
Novacek, M. J. (1991). “All tree histograms” and the evaluation of cladistic evidence: Some ambiguities. Cladistics 7: 345–349.
Novacek, M. J. (1992). Mammal phylogeny: Shaking the tree. Nature 356: 121–125.
Novacek, M. J., Rougier, G. W., Wible, J. R., McKenna, M. C., Dashzeveg, D., and Horovitz, I. (1997). Epipubic bones in eutherian mammals from the late Cretaceous of Mongolia. Nature 389: 483–486.
Novacek, M. J., and Wyss, A. R. (1986). Higher level relationships of the recent eutherian orders: The morphological evidence. Cladistics 2: 257–287.
O'Leary, M. O., and J. H. Geisler. (1999). The position of Cetace' within Mammlia: Phylogenetic analysis of morphological data from extinct and extant taxa. Syst. Biol. 48: 455–490.
Pol, D., and Siddall, M. E. (2001). Biases in maximum likelihood and parsimony: A simulation approach to a 10-taxon case. Cladistics 17: 266–281.
Radinsky, L. B. (1969). The early evolution of the Perissodactyla. Evolution 23: 308–328.
Reinhart, R. H. (1959). A review of the Sirenia and Desmostylia. Univ. Calif. Pub. Geol. Sci. 36: 1–146.
Renfree, M. B. (1993). Ontogeny, fenetic control, and phylogeny of female reproduction in monotreme and therian mammals. In: Mammal Phylogeny, Vol. 1: Mes. Diff, Multi, Mono, Early Therians, and Mammals, F. S. Szalay, M. J. Novacek, and M. C. McKenna, eds., pp. 4–20, Springer-Verlag, New York.
Rose, K. D., and Emry, R. J. (1993). Relationships of Xenarthra, Pholidota and fossil “edentates”: The morphological evidence. In: Mammal Phylogeny, Vol. 2: Placentals, F. S. Szalay, M. J. Novacek, and M. C. McKenna, eds., pp. 81–102, Springer-Verlag, New York.
Ross, C. F. (1995). Muscular and osseous anatomy of the primate anterior temporal fossa and the functions of the postorbital septum. Amer. J. Phys. Anthropol. 98: 275–306.
Ross, C. F., Williams, B., and Kay, R. F. (1998). Phylogenetic analysis of anthropoid relationships. J. Hum. Evol. 35: 221–306.
Schulmeister, S., Wheeler, W. C., and Carpenter, J. M. (2002). Simultaneous analysis of the basal lineages of Hymenoptera (Insecta) using sensitivity analysis. Cladistics 18: 455–484.
Schutt, W. A., and Simmons, N. B. (1998). Morphology and homology of the chiropteran calcar, with comments on the phylogenetic relationships of Archaeopteropus. J. Mammal. Evol. 5: 1–32.
Shikama, T. (1966). Postcranial skeletons of Japanese demostylians. Paleontol. Soc. Jpn. Sp. Paper 12: 1–202.
Shoshani, J., and McKenna, M. C. (1998). Higher taxonomic relationships among extant mammals based on morphology, with selected comparisons of results from molecular data. Mol. Phylogenet. Evol. 9: 572–584.
Shoshani, J., West, R. M., Court, N., Savage, R. J. G., and Harris, J. M. (1996). The earliest proboscideans: General plan, taxonomy, and palaeoecology. In: The Proboscidea: Evolution and Paleoecology of Elephants and Their Relatives, J., Shoshani and P. Tassy, eds., pp. 57–75, Oxford University Press, Oxford.
Siddall, M. E. (1998). Success of parsimony in the four-taxon case: Long-branch repulsion by likelihood in the Farris zone. Cladistics 14: 209–220.
Siddall, M. E., and Whiting, M. F. (1999). Long branch abstractions. Cladistics 15: 9–24.
Simmons, N. B. (1994). The case for chiropteran monophyly. Amer. Mus. Novit. 3103: 1–54.
Simmons, N. B. (1995). Bat reltionships and the origin of flight. Symp. Zool. Soc. Lond. 67: 27–43.
Simmons, N. B., and Geisler, J. H. (1998). Phylogenetic relationships of Icaronycteris, Archaeonycteris, Hassianycteris, and Palaeochiropteryx to extant bat lineages, with comments of the evolution of echolocation and foraging strategies in Microchiroptera. Bull. Amer. Mus. Nat. Hist. 235: 1–182.
Simons., E. L., and Rasmussen, D. T. (1989). Cranial morphology of Aegyptopithecus and Tarsius and the question of the tarsier-anthropoidean clade. Amer. J. Phys. Anthropol., 79: 1–23.
Simpson, G. G. (1931). A new insectivore from the Oligocene, Ulan Gochu horizon of Mongolia. Amer. Mus. Novit. 505: 1.
Simpson, G. G. (1945). The principles of classification and a classification of mammals. Bull. Amer. Mus. Nat. Hist. 85: 1–350.
Smith, A. B. (1998). What does paleontology contribute to systematics in a molecular world? Mol. Phylogenet. Evol. 9: 437–447.
Springer, M. S., Murphy, W. J., Eizirik, E., and O'Brien, S. J. (2003). Placental mammal diversification and the Cretaceous Tertiary boundary. Proc. Natl. Acad. Sci. U.S.A. 100: 1056–1061.
Stanhope, M. J., Waddell, V. G., Madsen, O., de Jong, W. W., Hedges, S. B., Cleven, G. C., Kao, D., and Springer, M.S. (1998). Molecular evidence for multiple origins of the Insectivora and for a new order of endemic African mammals. Proc. Natl. Acad. Sci. U.S.A. 95: 9967–9972.
Starck, D. (1995). Lehrbuch der Speziellen Zoologie; Band II: Wirbeltiere; Teile 1–2, Saeugetiere, Gustav Fischer Verlag, Jena.
Swofford, D. L. (2000). Phylogenetic Analysis Using Parsimony (*and Other Methods), Version 4.0b10, Sinauer Associates, Sunderland, MA.
Swofford, D. L., Waddell, P. J., Huelsenbeck, J. P., Foster, P. G., Lewis, P. O., and Rogers, J. S. (2001). Bias in phylogenetic estimation and its relevance to the choice between parsimony and likelihood methods. Syst. Biol. 50: 525–539.
Szalay, F. S., and Lucas, S. (1993). Cranioskeletal morphology of archontans, and diagnoses of Chiroptera, Volitantia, and Archonta. In: Primates and Their Relatives in Phylogenetic Perspective, R. D. E. MacPhee, ed., pp. 187–226, Plenum, New York.
Tabuce, R., Coiffait, B., Coiffait, P.-E., Mahboubi, M., and Jaeger, J.-J. (2001). A new genus of Macroscelidea (Mammalia) from the Eocene of Algeria: A possible origin of elephant shrews. J. Vertebr. Paleontol. 21: 535–546.
Tassy, P. (1981). Le crane de Moeritherium (Proboscidea, Mammalia) de l'Eocene de Dor el Talha (Libye) et le probleme de la classification phylogenetique du genre dans les Tethytheria. Bull. Mus Natn. Hist. Nat. 3: 87–147.
Thewissen, J. G. M. (1989). Mammlian frontal diploic vein and the human foramen caecum. Anat. Rec. 223: 242–244.
Thewissen, J. G. M. (1990). Evolution of Paleocene and Eocene Phenacodontidae (Mammalia, Condylarthra). Univ. Mich. Pap. Paleontol. 29: 1–107.
Thewissen, J. G. M. (1994). Phylogenetic aspects of cetacean origins: A morpholoical perspective. J. Mammal. Evol. 2: 157–184.
Thewissen, J. G. M., and Domning, D. (1992). The role of phenacodontids in the origin of the modern orders of ungulate mammals. J. Vertebr. Paleontol. 12: 494.
Thompson, J. D., Gibson, T. J., Plewniak, F., Jeanmougin, F., and Higgins, D. G. (1997). The CLUSTAL-X windows interface: Flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleie. Acids Res. 25: 4876–4882.
True, F. W. (1904). The whalebone whales of the western North Atlantic. Smithson. Contrib. Knowledge 33: 1–332.
Vanderhoof, V. L. (1937). A study of the Miocene sirenian Desmostylus. Univ. Calif. Pubs. Geol. Sci. 24: 119–262.
Van der Klaauw, C. J. (1931). The auditory bulla in some fossil mammals, with a general introduction to this region of the skull. Bull. Amer. Mus. Nat. Hist. 62: 1–352.
van Dijk, M. A. M, Paradis, E., Catzeflis, F., and de Jong, W. W. (1999). The virtues of gaps: Xenarthran (edentate) monophyly supported by a unique deletion in A-crystallin. Syst. Biol. 48: 94–106.
Webb, S. D., and Taylor, B. E. (1980). The phylogeny of hornless ruminants and a description of the cranium of Archaeomeryx. Bull. Amer. Mus. Nat. Hist. 167: 121–157.
Werdelin, L., and Nilsonne, A. (1999). The evolution of the scrotum and testicular descent in mammals: A phylogenetic view. J. Theor. Biol. 196: 61–72.
Wheeler, W. C. (1995). Sequence alignment, parameter sensitivity, and the phylogenetic analysis of molecular data. Syst. Biol. 44: 321–331.
Wheeler, W. C. (1996). Optimization alignment: The end of multiple sequence alignment in phylogenetics? Cladistics 12: 1–9.
Wheeler, W. C. (1999). Measuring topological congruence by extending character techniques Cladistics 15: 131–136.
Wheeler, W. C. (2001). Homology and the optimization of DNA sequence data. Cladistics 17: S3-S11.
Wheeler, W. C., Gladstein, D., and DeLaet, J. (2002). POY direct optimzation computer program, Versions test_5 through 3.0.5. Available at ftp.amnh.org/pub/molecular/poy.
Wible, J. R., and Novacek, M. J. (1988). Cranial evidence for the monophyletic origin of bats. Amer. Mus. Novit. 2911: 1–19.
Wible, J. R., Novacek, M. J., and Rougier, G. W. (in press). New data on the skull structure in the Mongolian Late Cretaceous eutherian mammal Zalambdalestes. Bull. Amer. Mus. Nat. Hist.
Wible, J. R., and Rougier, G. W. (2000). Cranial anatomy of Kryptobataar dashzevegi (Mammlia, Multituberculata) and its bearing on the evolution of mammalian characters. Bull. Amer. Mus. Nat. Hist. 247: 1–124.
Wible, J. R., Rougier, G. W., Novacek, M. J., and McKenna, M. C. (2001). Earliest eutherian ear region: A petrosal referred to Prokennalestes from the Early Cretaceous of Mongolia. Amer. Mus. Novit. 3322: 1–44.
Williamson, T. E., and Lucas, S. G. (1992). Meniscotherium (Mammalia, “Condylarthra”) from the Paleocene–Eocene of western North America. New Mex. Mus. Nat. Hist. Bull. 1: 1–75.
Wyss, A. D. (1987). Notes on Proteutheria, Insectivora, and Thomas Huxley's contribution to mammalian systematics. J. Mammal. 68: 135–138.
Wyss, A. R., and J. J. Flynn. (1993). A phylogenetic analysis and definition of the Carnivora. In: Mammal Phylogeny, Vol. 2: Placentals, F. S. Szalay, M. J. Novacek, and M. C. McKenna, eds., pp. 32–52, Springer-Verlag, New York.
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Asher, R.J., Novacek, M.J. & Geisler, J.H. Relationships of Endemic African Mammals and Their Fossil Relatives Based on Morphological and Molecular Evidence. Journal of Mammalian Evolution 10, 131–194 (2003). https://doi.org/10.1023/A:1025504124129
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DOI: https://doi.org/10.1023/A:1025504124129