Abstract
All materials, including upper and lower jaw fragments and isolated teeth, of two closely related species of the stem placental mammal Prokennalestes, P. minor Kielan-Jaworowska et Dashzeveg, 1989 (258 specimens) and P. trofimovi Kielan-Jaworowska et Dashzeveg, 1989 (251 specimens), from the type locality of Khovoor (Early Cretaceous of Mongolia) are described. These extensive materials allow for the first time the study of morphological and size variability in an Early Cretaceous mammal. Prokennalestes can be diagnosed by the dental formula I?/4, C1/1, P4–5/5, M3/3, presence of the Meckelian groove, the mandibular foramen positioned dorsally on a prominent longitudinal ridge, presence of the masseteric foramen, double-rooted and premolariform lower canine, submolariform P5 with a large protocone and minute metacone, presence of the preparastyle, cusplike postmetacrista, and cusplike conules with incipient conular cristae on the upper molars, premolariform p5, the protoconid distinctly higher than the metaconid, the paraconid and metaconid connate at the base, transverse protocristid, and presence of distal metacristid on the lower molars. Two Prokennalestes species are reliably distinguished by the tooth dimensions (P. minor is smaller by 20%), the position of the infraorbital foramen (under P4 in P. minor and between P4 and P5 or under P5 in P. trofimovi), the presence of protoconal swelling on P4 (usually absent in P. minor and usually present in P. trofimovi), the ectocingulum extension in M3 (longer in P. minor), position of the posterior end of the mandibular symphysis (under p2 in P. minor, under p2–p3 in P. trofimovi), and by the position of the posterior mental foramen (under p4 in P. minor and under p5 in P. trofimovi). Taking into account considerable morphological differences between the two samples, they are regarded as different species rather than sexual morphs of the same species. Prokennalestes is a basal eutherian mammal similar in phylogenetic position to other stem placentals (Eomaia, Murtoilestes, Bobolestes, Aspanlestes). A more exact phylogenetic position of Prokennalestes can be established based on rigorous phylogenetic analysis including a large set of known stem therian and stem placental taxa.
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References
Archibald, J.D., A study of Mammalia and geology across the Cretaceous–Tertiary boundary in Garfield County, Montana, Univ. California Publ. Geol. Sci., 1982, vol. 122, pp. 1–286.
Archibald, J.D. and Averianov, A.O., The Late Cretaceous placental mammal Kulbeckia, J. Vertebr. Paleontol., 2003, vol. 23, no. 2, pp. 404–419.
Archibald, J.D. and Averianov, A.O., Late Cretaceous asioryctitherian eutherian mammals from Uzbekistan and phylogenetic analysis of Asioryctitheria, Acta Palaeontol. Polon., 2006, vol. 51, no. 2, pp. 351–376.
Archibald, J.D. and Averianov, A.O., Phylogenetic analysis, taxonomic revision, and dental ontogeny of the Cretaceous Zhelestidae (Mammalia, pp. Eutheria), Zool. J. Linnean Soc., 2012, vol. 164, no. 2, pp. 361–426.
Averianov, A.O., Early Cretaceous “symmetrodont” mammal Gobiotheriodon from Mongolia and the classification of “Symmetrodonta”, Acta Palaeontol. Polon., 2002, vol. 47, no. 4, pp. 705–716.
Averianov, A.O., Taxonomic revision of tribosphenic mammals from the Lower Cretaceous Antlers Formation of Texas and Oklahoma, USA, Proc. Zool. Inst. Russ. Acad. Sci., 2015, vol. 319, no. 2, pp. 141–181.
Averianov, A.O. and Archibald, J.D., Mammals from the mid–Cretaceous Khodzhakul Formation, Kyzylkum Desert, Uzbekistan, Cret. Res., 2005. 26, no. 4, pp. 593–608.
Averianov, A.O. and Archibald, J.D., New material and reinterpretation of the Late Cretaceous eutherian mammal Paranyctoides from Uzbekistan, Acta Palaeontol. Polon., 2013a, vol. 58, no. 1, pp. 17–23.
Averianov, A.O., and Archibald, J.D., Variation and taxonomy of Asiamerican eutherian mammal Paranyctoides, Can. J. Earth Sci., 2013b, vol. 50, no. 9, pp. 895–903.
Averianov, A.O. and Archibald, J.D., Evolutionary transition of dental formula in Late Cretaceous eutherian mammals, Sci. Nature, 2015, vol. 102, nos. 9–10, p. 56.
Averianov, A.O. and Archibald, J.D., New evidence on the stem placental mammal Paranyctoides from the Upper Cretaceous of Uzbekistan, Palaeontol. Polon., 2016, vol. 67, pp. 25–33.
Averianov, A.O., Archibald, J.D., and Dyke, G.J., A new eutherian mammal from the Late Cretaceous of Kazakhstan, Acta Palaeontol. Polon., 2014a, vol. 59, no. 3, pp. 537–542.
Averianov, A.O., Martin, T., and Lopatin, A.V., A new phylogeny for basal Trechnotheria and Cladotheria and affinities of South American endemic Late Cretaceous mammals, Naturwissenschaften, 2013, vol. 100, no. 4, pp. 311–326.
Averianov, A.O., Martin, T., and Lopatin, A.V., The oldest dryolestid mammal from the Middle Jurassic of Siberia, J. Vertebr. Paleontol., 2014b, vol. 34, no. 4, pp. 924–931.
Averianov, A.O., Martin, T., Lopatin, A.V., and Krasnolutskii, S.A., Stem therian mammal Amphibetulimus from the Middle Jurassic of Siberia, Paläontol. Z., 2015, vol. 89, no. 2, pp. 197–206.
Averianov, A.O. and Skutschas, P.P., Phylogenetic relationships within basal tribosphenic mammals, Proc. Zool. Inst. Russ. Acad. Sci., 1999, vol. 281, pp. 55–60.
Averianov, A.O. and Skutschas, P.P., An eutherian mammal from the Early Cretaceous of Russia and biostratigraphy of the Asian Early Cretaceous vertebrate assemblages, Lethaia, 2000, vol. 33, no. 4, pp. 330–340.
Averianov, A.O. and Skutschas, P.P., A new genus of eutherian mammal from the Early Cretaceous of Transbaikalia, Russia, Acta Palaeontol. Polon., 2001, vol. 46, no. 3, pp. 431–436.
Barsbold, R., Voronin, Y.L., and Zhegallo, V.I., On the work of Soviet–Mongolian Paleontological Expedition in 1969–1970 years, Paleontologicheskii Zh., 1971, no. 2, pp. 139–143.
Beck, R.M.D. and Lee, M.S.Y., Ancient dates or accelerated rates? Morphological clocks and the antiquity of placental mammals, Proc. Royal Soc. B Biol. Sci., 2014, vol. 281, no. 1793, p. 20141278.
Belyaeva, E.I., Trofimov, B.A., and Reshetov, V.Y., General stages in evolution of late Mesozoic and early Tertiary mammalian faunas in Central Asia, Tr. Sovm. Sovet.-Mongol. Paleontol. Eksped., 1974, vol. 1, pp. 19–45.
Bi, S., Wang, Y., Guan, J., Sheng, X., and Meng, J., Three new Jurassic euharamiyidan species reinforce early divergence of mammals, Nature, 2014, vol. 514, no. 7524, pp. 579–584.
Bi, S., Zheng, X., Meng, J., Wang, X., Robinson, N., and Davis, B., A new symmetrodont mammal (Trechnotheria, pp. Zhangheotheriidae) from the Early Cretaceous of China and trechnotherian character evolution, Sci. Rep., 2016, vol. 6, p. 26668.
Bonaparte, J.F. and Rougier, G.W., Mamiferos del Cretacico Inferior de Patagonia, Argentina, in IV Congreso Latinamericano de Paleontologia, 1987, vol. 1, pp. 343–359.
Butler, P.M. and Clemens, W.A., Dental morphology of the Jurassic holotherian mammal Amphitherium, with a discussion of the evolution of mammalian postcanine dental formulae, Palaeontology, 2001, vol. 44, no. 1, pp. 1–20.
Cifelli, R.L. and Madsen, S.K., Spalacotheriid symmetrodonts (Mammalia) from the medial Cretaceous (upper Albian or lower Cenomanian) Mussentuchit local fauna, Cedar Mountain Formation, Utah, USA, Geodiversitas, 1999, vol. 21, no. 2, pp. 167–214.
Clemens, W.A., Fossil mammals of the type Lance Formation, Wyoming: Part III. Eutheria and summary, Univ. California Publ. Geol. Sci., 1973, vol. 94, pp. 1–102.
Clemens, W.A. and Lillegraven, J.A., New Late Cretaceous, North American advanced therian mammals that fit neither the marsupial nor eutherian molds, Contrib. Geol. Univ. Wyoming Spec. Pap., 1986, vol. 3 (Vertebrates, Phylogeny and Philosophy, Flanagan, K.M. and Lillegraven, J.A., Eds.), pp. 55–86.
Clemens, W.A. and Mills, J.R.E., Review of Peramus tenuirostris Owen (Eupantotheria, Mammalia), Bull. Brit. Mus. Natur. Hist. Geol., 1971, vol. 20, pp. 89–113.
Close, R.A., Davis, B.M., Walsh, S.A., Wolniewicz, A.S., Friedman, M., and Benson, R.B.J., A lower jaw of Palaeoxonodon from the Middle Jurassic of the Isle of Skye, Scotland, sheds new light on the diversity of British stem therians, Palaeontology, 2016, vol. 59, no. 1, pp. 155–169.
Cohen, J.E., Earliest divergence of stagodontid (Mammalia: Marsupialiformes) feeding strategies from the Late Cretaceous (Turonian) of North America, J. Mamm. Evol., 2017, doi: 10.1007/s10914-017-9382-0
Crompton, A.W., The origin of the tribosphenic molar, Zool. J. the Linnean Soc., 1971, vol. 50 (Early Mammals, Kermack, D.M. and Kermack, K.A., Eds.), suppl. no. 1, pp. 65–87.
Crompton, A.W. and Kielan-Jaworowska, Z., Molar structure and occlusion in Cretaceous therian mammals, in Studies in the Development, Function and Evolution of Teeth, Butler P.M. and Joysey, K.A., Eds., London: Acad. Press, 1978, pp. 249–287.
Dashzeveg, D. and Kielan-Jaworowska, Z., The lower jaw of an aegialodontid mammal from the Early Cretaceous of Mongolia, Zool. J. Linnean Soc., 1984, vol. 82, pp. 217–227.
Dashzeveg, D., Reshetov, V.Y., and Trofimov, B.A., Early stages of development of mammals in Mongolia, in Main Results of Investigations of the Joint Soviet–Mongol Paleontological Expedition for 1969–1988 years. Abstracts of Reports, Moscow: Paleontol. Inst. Akad. Nauk SSSR, 1989, pp. 5–6.
Davis, B.M., Evolution of the tribosphenic molar pattern in early mammals, with comments on the “dual–origin” hypothesis, J. Mammal. Evol., 2011, vol. 18, no. 4, pp. 227–244.
Davis, B.M., Micro-computed tomography reveals a diversity of peramuran mammals from the Purbeck Group (Berriasian) of England, Palaeontology, 2012, vol. 55, no. 4, pp. 789–817.
Davis, B.M., Cifelli, R.L., and Kielan-Jaworowska, Z., Earliest evidence of Deltatheroida (Mammalia: Metatheria) from the Early Cretaceous of North America, in Mammalian Evolutionary Morphology. A Tribute to Frederick S. Szalay, Sargis, E.J. and Dagosto, M., Eds., Springer, 2008, pp. 3–24.
Ekdale, E.G., Archibald, J.D., and Averianov, A.O., Petrosal bones of placental mammals from the Late Cretaceous of Uzbekistan, Acta Palaeontol. Polon., 2004, vol. 49, no. 1, pp. 161–176.
Fostowicz-Frelik, L., Convergent and parallel evolution in early Glires (Mammalia), in Evolutionary Biology: Self/Nonself Evolution, Species and Complex Traits Evolution, Methods and Concepts, Pontarotti, P., Ed., Springer, 2017, pp. 199–216.
Fox, R.C., Molar structure and function in the Early Cretaceous mammal Pappotherium: Evolutionary implications for Mesozoic Theria, Can. J. Earth Sci., 1975, vol. 12, no. 3, pp. 412–442.
Fox, R.C., Notes on the dentition and relationships of the Late Cretaceous insectivore Gypsonictops, Can. J. Earth Sci., 1977, vol. 14, no. 8, pp. 1823–1831.
Gheerbrant, E., Bustylus (Eutheria, Adapisoriculidae) and absence of ascertained marsupials in the Palaeocene of Europe, Terra Nova, 1991, vol. 3, no. 6, pp. 586–592.
Gill, T., Arrangement of the families of mammals. With analytical tables, Smithsonian Miscellan. Coll., 1872, vol. 11, no. 230, pp. 1–98.
Goswami, A., Prasad, G.V.R., Upchurch, P., Boyer, D.M., Seiffert, E.R., Verma, O., Gheerbrant, E., and Flynn, J.J., A radiation of arboreal basal eutherian mammals beginning in the Late Cretaceous of India, Proc. Nat. Acad. Sci. USA, 2011, vol. 27, no. 108, pp. 16333–16338.
Han, G. and Meng, J., A new spalacolestine mammal from the Early Cretaceous Jehol Biota and implications for the morphology, phylogeny, and palaeobiology of Laurasian ‘symmetrodontans’, Zool. J. Linnean Soc., 2016, vol. 178, no. 2, pp. 343–380.
Hu, Y., Fox, R.C., Wang, Y., and Li, C., A new spalacotheriid symmetrodont from the Early Cretaceous of Northeastern China, Am. Mus. Novit., 2005, vol. 3475, pp. 1–20.
Hu, Y., Meng, J., Li, C., and Wang, Y., New basal eutherian mammal from the Early Cretaceous Jehol biota, Liaoning, China, Proc. Royal Soc. B Biol. Sci., 2010, vol. 277, no. 1679, pp. 229–236.
Jerzykiewicz, T. and Russell, D.A., Late Mesozoic stratigraphy and vertebrates of the Gobi Basin, Cret. Res., 1991, vol. 12, pp. 345–377.
Ji, Q., Luo, Z.-X., and Ji, S.-a., A Chinese triconodont mammal and mosaic evolution of mammalian skeleton, Nature, 1999, vol. 398, no. 6725, pp. 326–330.
Ji, Q., Luo, Z.-X., Yuan, C.-X., Wible, J.R., Zhang, J.-P., and Georg, J.A., The earliest known eutherian mammal, Nature, 2002, vol. 416, no. 6883, pp. 816–822.
Ji, Q., Luo, Z.-X., Zhang, X., Yuan, C.-X., and Xu, L., Evolutionary development of the middle ear in Mesozoic therian mammals, Science, 2009, vol. 326, no. 5950, pp. 278–281.
Kalandadze, N.N. and Kurzanov, S.M., Lower Cretaceous localities of terrestrial vertebrates in Mongolia, Tr. Sovm. Sovet.-Mongol. Paleontol. Eksped., 1974, vol. 1, pp. 288–295.
Kalandadze, N.N. and Reshetov, V.Y., Interesting paleontological findings in Mongolia, Priroda, 1971, no. 5, pp. 83–84.
Kemp, T.S., The Origin and Evolution of Mammals, Oxford–New York: Oxford Univ. Press, 2005.
Kermack, K.A., Lees, P.M., and Mussett, F., Aegialodon dawsoni, a new trituberculosectorial tooth from Lower Wealden, Proc. Royal Soc. London Ser. B Biol. Sci., 1965, vol. 162, no. 989, pp. 535–554.
Kielan-Jaworowska, Z., Evolution of the therian mammals in the Late Cretaceous of Asia. Part III. Postcranial skeleton in Zalambdalestidae, Palaeontol. Polon., 1978, vol. 38, pp. 5–41.
Kielan-Jaworowska, Z., Evolution of the therian mammals in the Late Cretaceous of Asia: Part IV. Skull structure in Kennalestes and Asioryctes, Palaeontol. Polon., 1981, vol. 42, pp. 25–78.
Kielan-Jaworowska, Z., Evolution of the therian mammals in the Late Cretaceous of Asia: Part VII. Synopsis. Palaeontol. Polon., 1984a, vol. 46, pp. 173–183.
Kielan-Jaworowska, Z., Evolution of the therian mammals in the Late Cretaceous of Asia: Part V. Skull structure in Zalambdalestidae, Palaeontol. Polon., 1984b, vol. 46, pp. 107–117.
Kielan-Jaworowska, Z., Bown, T.M., and Lillegraven, J.A., Eutheria, in Mesozoic Mammals: The First Two–Thirds of Mammalian History, Lillegraven, J.A., Kielan-Jaworowska., and Clemens, W.A., Eds., Berkeley: Univ. California Press, 1979, pp. 221–258.
Kielan-Jaworowska, Z., Cifelli, R.L., and Luo, Z.-X., Alleged Cretaceous placental from down under, Lethaia, 1998, vol. 31, no. 3, pp. 267–268.
Kielan-Jaworowska, Z., Cifelli, R.L., and Luo, Z.-X., Mammals from the Age of Dinosaurs, pp. Origins, Evolution, and Structure, New York, Columbia Univ. Press, 2004.
Kielan-Jaworowska, Z. and Dashzeveg, D., Eutherian mammals from the Early Cretaceous of Mongolia, Zool. Scr., 1989, vol. 18, pp. 347–355.
Kielan-Jaworowska, Z., Novacek, M.J., Trofimov, B.A., and Dashzeveg, D., Mammals from the Mesozoic of Mongolia, in The Age of Dinosaurs in Russia and Mongolia, Benton, M.J., Shishkin, M.A., Unwin, D.M., and Kurochkin, E.N., Eds., Cambridge: Cambridge Univ. Press, 2000, pp. 573–626.
Kielan-Jaworowska, Z. and Trofimov, B.A., A new occurrence of Late Cretaceous eutherian mammal Zalambdalestes, Acta Palaeontol. Polon., 1981, vol. 26, pp. 3–7.
Kramarenko, N.N., On the work of the Joint Soviet–Mongolian Paleontological Expedition during the years 1969–1972, Tr. Sovm. Sovet.–Mongol. Paleontol. Eksped., 1974, vol. 1, pp. 9–18.
Krassilov, V.A., Early Cretaceous flora of Mongolia, Palaeontogr. Abt. B Palaeobot., 1982, vol. 181, pp. 1–43.
Krebs, B., Das Skelett von Henkelotherium guimarotae gen. et sp. nov. (Eupantotheria, Mammalia) aus dem Oberen Jura von Portugal, Berl. Geowiss. Abh., Reihe A Geol. Palaeontol., 1991, vol. 133, pp. 1–121.
Kurochkin, E.N., Kalandadze, N.N., and Reshetov, V.Y., First results of the Soviet–Mongolian Paleontological Expedition, Priroda, 1970, no. 4, p. 115.
Kusuhashi, N., Tsutsumi, Y., Saegusa, H., Horie, K., Ikeda, T., Yokoyama, K., and Shiraishi, K., A new Early Cretaceous eutherian mammal from the Sasayama Group, Hyogo, Japan, Proc. Royal Soc. B Biol. Sci., 2013, vol. 280, no. 1759, p. 20130142.
Li, G. and Luo, Z.-X., A Cretaceous symmetrodont therian with some monotreme-like postcranial features, Nature, 2006, vol. 439, no. 7073, pp. 195–200.
Lillegraven, J.A., 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., 1969, vol. 50, pp. 1–122.
Linnaeus, C., Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis, vol. 1: Regnum animale. Editio decima, reformata, Stockholm: Laurentius Salvius, 1758.
Lopatin, A.V. and Averianov, A.O., Revision of a pretribosphenic mammal Arguimus from the Early Cretaceous of Mongolia, Acta Palaeontol. Polon., 2006, vol. 51, no. 2, pp. 339–349.
Lopatin, A.V. and Averianov, A.O., Kielantherium, a basal tribosphenic mammal from the Early Cretaceous of Mongolia, with new data on the aegialodontid dentition, Acta Palaeontol. Polon., 2007, vol. 52, no. 3, pp. 441–446.
Lopatin, A.V. and Averianov, A.O., Early Cretaceous mammals from the Khoboor in Mongolia, in Paleontology of Central Asia: Abstracts of the International Conference on the 40th Anniversary of the Joint Russian–Mongolian Paleontological Expedition. Moscow, November 18–19, 2009, Moscow, 2009, pp. 57–60.
Lopatin, A.V. and Averianov, A.O., Gobiconodon (Mammalia) from the Early Cretaceous of Mongolia and revision of Gobiconodontidae, J. Mammal. Evol., 2015, vol. 22, pp. 17–43.
Lopatin, A.V., Averianov, A.O., Maschenko, E.N., and Leshchinskiy, S.V., Early Cretaceous mammals of Western Siberia: 3. Zhangheotheriidae, Paleontol. J., 2010, vol. 44, no. 5, pp. 573–583.
Lopatin, A.V., Maschenko, E.N., Averianov, A.O., Rezvyi, A.S., Skutschas, P.P., and Leshchinskiy, S.V., Early Cretaceous mammals from Western Siberia: 1. Tinodontidae, Paleontol. J., 2005, vol. 39, no. 5, pp. 523–534.
Lucas, S.G., The Psittacosaurus biochron, Early Cretaceous of Asia, Cret. Res., 2006, vol. 27, no. 2, pp. 189–198.
Luo, Z.-X., Chen, P., Li, G., and Chen, M., A new eutriconodont mammal and evolutionary development in early mammals, Nature, 2007a, vol. 446, no. 7133, pp. 288–293.
Luo, Z.-X. and Ji, Q., New study on dental and skeletal features of the Cretaceous “symmetrodontan” mammal Zhangheotherium, J. Mammal. Evol., 2005, vol. 12, nos. 3/4, pp. 337–357.
Luo, Z.-X., Ji, Q., Wible, J.R., and Yuan, C.-X., An Early Cretaceous tribosphenic mammal and metatherian evolution, Science, 2003, vol. 302, no. 5652, pp. 1934–1940.
Luo, Z.-X., Ji, Q., and Yuan, C.-X., Convergent dental adaptations in pseudo–tribosphenic and tribosphenic mammals, Nature, 2007b, vol. 450, no. 7166, pp. 93–97.
Luo, Z.-X., Kielan-Jaworowska, Z., and Cifelli, R.L., In quest for a phylogeny of Mesozoic mammals, Acta Palaeontol. Polon., 2002, vol. 47, no. 1, pp. 1–78.
Luo, Z.-X., Meng, Q.-J., Ji, Q., Liu, D., Zhang, Y.-G., and Neander, A.I., Evolutionary development in basal mammaliaforms as revealed by a docodontan, Science, 2015, vol. 347, no. 6223, pp. 760–764.
Luo, Z.-X. and Wible, J.R., A Late Jurassic digging mammal and early mammalian diversification, Science, 2005, vol. 308, no. 5718, pp. 103–107.
Luo, Z.-X., Yuan, C.-X., Meng, Q.-J., and Ji, Q., A Jurassic eutherian mammal and divergence of marsupials and placentals, Nature, 2011, vol. 476, no. 7361, pp. 442–445.
Manz, C.L., Chester, S.G.B., Bloch, J.I., Silcox, M.T., and Sargis, E.J., New partial skeletons of Palaeocene Nyctitheriidae and evaluation of proposed euarchontan affinities, Biol. Lett., 2015, vol. 11, no. 1.
Martin, T., Dryolestidae (Dryolestoidea, Mammalia) aus dem Oberen Jura von Portugal, Abh. Senckenberg. Naturforsch. Gesell., 1999, vol. 550, pp. 1–119.
Martin, T., New stem-line representatives of Zatheria (Mammalia) from the Late Jurassic of Portugal, J. Vertebr. Paleontol., 2002, vol. 22, no. 2, pp. 332–348.
McKenna, M.C., Towards a phylogenetic classification of the Mammalia, in Phylogeny of the Primates, Luckett, W.P. and Szalay, F.S., New York: Plenum Press, 1975, pp. 21–46.
McKenna, M.C. and Bell, S.K., Classification of Mammals above the Species Level, New York: Columbia Univ. Press, 1997.
Meng, J., Hu, Y., Wang, Y., Wang, X., and Li, C., A Mesozoic gliding mammal from northeastern China, Nature, 2006, vol. 444, no. 7121, pp. 889–893.
Meng, J., Wang, Y., and Li, C., Transitional mammalian middle ear from a new Cretaceous Jehol eutriconodont, Nature, 2011, vol. 472, no. 7342, pp. 181–185.
Montellano-Ballesteros, M., Fox, R.C., and Scott, C.S., Species composition of the Late Cretaceous eutherian mammal Paranyctoides Fox, Can. J. Earth Sci., 2013, vol. 50, no. 7, pp. 693–700.
Muizon, C., Billet, G., Argot, C., Ladevéze, S., and Goussard, F., Alcidedorbignya inopinata, a basal pantodont (Placentalia, Mammalia) from the early Palaeocene of Bolivia: Anatomy, phylogeny and palaeobiology, Geodiversitas, 2015, vol. 37, no. 4, pp. 397–634.
Nesov, L.A., New mammals from the Cretaceous of Kyzylkum, Vest. Leningr. Univ., Ser. 7, 1985, no. 17, pp. 8–18.
Nesov, L.A., Archibald, J.D., and Kielan-Jaworowska, Z., Ungulate-like mammals from the Late Cretaceous of Uzbekistan and a phylogenetic analysis of Ungulatomorpha, Bull. Carnegie Mus. Natur. Hist., 1998, vol. 34, pp. 40–88.
Ni, X., Li, Q., Stidham, T.A., Li, L., Lu, X., and Meng, J., A late Paleocene probable metatherian (?deltatheroidan) survivor of the Cretaceous mass extinction, Sci. Rep., 2016, vol. 6, p. 38547.
Novacek, M.J., The primitive eutherian dental formula, J. Vertebr. Paleontol., 1986a, vol. 6, no. 2, pp. 191–196.
Novacek, M.J., The skull of leptictid insectivorans and the higher–level classification of eutherian mammals, Bull. Am. Mus. Natur. Hist., 1986b, vol. 183, pp. 1–112.
Novacek, M.J., Rougier, G.W., Wible, J.R., McKenna, M.C., Dashzeveg, D., and Horovitz, I., Epipubic bones in eutherian mammals from the Late Cretaceous of Mongolia, Nature, 1997, vol. 389, no. 6650, pp. 483–486.
Owen, R., Teeth, in The Cyclopedia of Anatomy and Physiology, Todd, R.B., Ed., London: Sherwood, Gilbert, and Piper, 1837, vol. 4, part 2, pp. 864–935.
Rasmussen, T.E. and Callison, G.L., A new species of triconodont mammal from the Upper Jurassic of Colorado, J. Paleontol., 1981, vol. 55, no. 3, pp. 628–634.
Reshetov, V.Y., Mammals of Central Asia, Priroda, 1989, no. 9, pp. 55–60.
Reshetov, V.Y. and Trofimov, B.A., The main stages of mammal development in Asia, in Paleontology. Stratigraphy. International Geological Congress, 26th Session, Sokolov, B.S., Ed., Moscow: Nauka, 1980, pp. 103–114.
Reshetov, V.Y. and Trofimov, B.A., Review of the study of fossil mammals from the USSR, in Theriology in the USSR, Sokolov, V.E. and Kucheruk, V.V., Eds., Moscow, 1984, pp. 6–29.
Rich, T.H.V., The palaeobiogeography of Mesozoic mammals: A review, Arqui. Mus. Nac., Rio de Janeiro, 2008, vol. 66, no. 1, pp. 231–249.
Rich, T.H.V., Flannery, T.F., Trusler, P., Kool, L., and Klaveren, N.A., and van Vickers-Rich, P., Evidence that monotremes and ausktribosphenids are not sister groups, J. Vertebr. Paleontol., 2002, vol. 22, no. 2, pp. 466–469.
Rich, T.H.V., Hopson, J.A., Gill, P.G., Trusler, P., Rogers-Davidson, S., Morton, S., Cifelli, R.L., Pickering, D., Kool, L., Siu, K., Burgmann, F.A., Senden, T., Evans, A.R., Wagstaff, B.E., Seegets-Villiers, D., Corfe, I.J., Flannery, T.F., Walker, K., Musser, A.M., Archer, M., Pian, R., and Vickers-Rich, P., The mandible and dentition of the Early Cretaceous monotreme Teinolophos trusleri, Alcheringa, 2016, vol. 40, no. 4, pp. 475–501.
Rougier, G.W., Apesteguia, S., and Gaetano, L.C., Highly specialized mammalian skulls from the Late Cretaceous of South America, Nature, 2011, vol. 479, no. 7371, pp. 98–102.
Rougier, G.W., Davis, B.M., and Novacek, M.J., A deltatheroidan mammal from the Upper Cretaceous Baynshiree Formation, eastern Mongolia, Cret. Res., 2015, vol. 52, pp. 167–177.
Rougier, G.W., Isaji, S., and Manabe, M., An Early Cretaceous mammal from the Kuwajima Formation (Tetori Group), Japan, and a reassessment of triconodont phylogeny, Ann. Carnegie Mus., 2007, vol. 76, no. 2, pp. 73–115.
Rougier, G.W., Wible, J.R., and Novacek, M.J., Implications of Deltatheridium specimens for early marsupial history, Nature, 1998, vol. 396, no. 6710, pp. 459–463.
Rougier, G.W., Wible, J.R., and Novacek, M.J., New specimen of Deltatheroides cretacicus (Metatheria, Deltatheroida) from the Late Cretaceous of Mongolia, Bull. Carnegie Mus. Natur. Hist., 2004, vol. 36, pp. 245–266.
Rowe, T.B., Rich, T.H.V., Vickers-Rich, P., Springer, M.S., and Woodburne, M.O., The oldest platypus and its bearing on divergence timing of the platypus and echidna clades, Proc. Nat. Acad. Sci., 2008, vol. 105, no. 4, pp. 1238–1242.
Shikama, T., Teilhardosaurus and Endotherium, new Jurassic Reptilia and Mammalia from the Husin coal-field, south Manchuria, Proc. Jap. Acad., 1947, vol. 23, pp. 76–84.
Shuvalov, V.F., On the geological structure and age of the localities Khoboor and Khuren–Dukh, Tr. Sovm. Sovet.- Mongol. Paleontol. Eksped., 1974, vol. 1, pp. 296–313.
Shuvalov, V.F., Stratigraphy of Mesozoic deposits of Cemtral Mongolia, Tr. Sovm. Sovet.-Mongol. Nauchno-Issled. Geol. Eksp., 1975, vol. 13, pp. 50–112.
Shuvalov, V.F., The Cretaceous stratigraphy and palaeobiogeography of Mongolia, in The Age of Dinosaurs in Russia and Mongolia, Benton, M.J., Shishkin, M.A., Unwin, D.M., and Kurochkin, E.N., Eds., Cambridge: Cambridge Univ. Press, 2000, pp. 256–278.
Sigogneau-Russell, D., Dashzeveg, D., and Russell, D.E., Further data on Prokennalestes (Mammalia, Eutheria, inc. sed.) from the Early Cretaceous of Mongolia, Zool.Scr., 1992, vol. 21, pp. 205–209.
Sigogneau-Russell, D., Hooker, J.J., and Ensom, P.C., The oldest tribosphenic mammal from Laurasia (Purbeck Limestone Group, Berriasian, Cretaceous, UK) and its bearing on the ‘dual origin’ of Tribosphenida, Comptes Rendus l’Acad. Sci., Ser. II, Fasc. A Sci. Terre Planètes, 2001, vol. 333, pp. 141–147.
Simpson, G.G., A Catalogue of the Mesozoic Mammalia in the Geological Department of the British Museum, London: British Mus. Natur. Hist., 1928.
Stucky, R.K. and McKenna, M.C., Mammalia, in The Fossil Record 2, Benton, M.J., Ed., London: Chapman and Hall, 1993, pp. 739–771.
Trofimov, B.A., Mammals from the Lower Cretaceous of Mongolia, in Session Devoted to the Hundred’s Anniversary of the Academichian A.A. Borissiak, Moscow: Paleontol. Inst. Akad. Nauk SSSR, 1972, pp. 65–67.
Trofimov, B.A. and Reshetov, V.Y., Asia as a center of mammals development, Priroda, 1975, no. 8, pp. 32–43.
Wible, J.R., On the cranial osteology of the short–tailed opossum Monodelphis brevicaudata (Didelphidae, Marsupialia), Ann. Carnegie Mus., 2003, vol. 72, no. 3, pp. 137–202.
Wible, J.R., Novacek, M.J., and Rougier, G.W., New data on the skull and dentition in the Mongolian Late Cretaceous eutherian mammal Zalambdalestes, Bull. American Museum of Natural Hist, 2004, vol. 281, pp. 1–144.
Wible, J.R., Rougier, G.W., and Novacek, M.J., Anatomical evidence for superordinal/ordinal eutherian taxa in the Cretaceous, in The Rise of Placental Mammals: Origins and Relationships of the Major Extant Clades, Rose, K.D. and Archibald, J.D., Eds., Baltimore: Johns Hopkins Univ. Press, 2005.
Wible, J.R., Rougier, G.W., Novacek, M.J., and Asher, R.J., Cretaceous eutherians and Laurasian origin for placental mammals near the K/T boundary, Nature, 2007, vol. 447, no. 7147, pp. 1003–1006.
Wible, J.R., Rougier, G.W., Novacek, M.J., and Asher, R.J., The eutherian mammal Maelestes gobiensis from the Late Cretaceous of Mongolia and the phylogeny of Cretaceous Eutheria, Bull. Am. Mus. Natur. Hist., 2009, vol. 327, pp. 1–123.
Wible, J.R., Rougier, G.W., Novacek, M.J., and McKenna, M.C., Earliest eutherian ear region: A petrosal referred to Prokennalestes from the Early Cretaceous of Mongolia, Am. Mus. Novit., 2001, vol. 3322, pp. 1–44.
Wilson, G.P., Ekdale, E.G., Hoganson, J.W., Calede, J.J., and Linden, A.V., A large carnivorous mammal from the Late Cretaceous and the North American origin of marsupials, Nature Commun., 2016, vol. 7, p. 13734.
Yabe, H. and Shikama, T., A new Jurassic mammal from south Manchuria, Proc. Imp. Acad. Tokyo, 1938, vol. 14, pp. 353–359.
Yuan, C.-X., Ji, Q., Meng, Q.-J., Tabrum, A.R., and Luo, Z.-X., Earliest evolution of multituberculate mammals revealed by a new Jurassic fossil, Science, 2013, vol. 341, no. 6147, pp. 779–783.
Zan, S., Wood, C.B., Rougier, G.W., Jin, L., Chen, J., and Schaff, C.R., A new “middle” Cretaceous zalambdalestid mammal, from a new locality in Jilin Province, northeastern China, J. Paleontol. Soc. Korea, 2006, vol. 22, no. 1, pp. 153–172.
Zheng, X., Bi, S., Wang, X., and Meng, J., A new arboreal haramiyid shows the diversity of crown mammals in the Jurassic period, Nature, 2013, vol. 500, no. 7461, pp. 199–202.
Zhou, C.-F., Wu, S., Martin, T., and Luo, Z.-X., A Jurassic mammaliaform and the earliest mammalian evolutionary adaptations, Nature, 2013, vol. 500, no. 7461, pp. 163–167.
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Lopatin, A.V., Averianov, A.O. The stem placental mammal Prokennalestes from the Early Cretaceous of Mongolia. Paleontol. J. 51, 1293–1374 (2017). https://doi.org/10.1134/S0031030117120048
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DOI: https://doi.org/10.1134/S0031030117120048