New Late Miocene Alilepus (Lagomorpha, Mammalia) from Eastern Europe - a new light on the evolution of the earliest Old World Leporinae


Authors: Čermák S, Angelone C, Sinitsa MV

Published in: Bulletin of Geosciences, volume 90, issue 2; pages: 431 - 451; Received 10 June 2014; Accepted in revised form 6 November 2014; Online 9 March 2015

Keywords: Alilepus, taxonomy, phylogeny, Late Miocene, Pliocene, Eastern Europe,

full text (PDF, 3.08 MB)

Export to RIS



The leporid material from eight Late Miocene (MN11-13) localities of Ukraine and Moldova is described. The analysis is supported by a direct revision of type materials of A. laskarewi, A. hungaricus and A. ucrainicus. Lectotypes of A. annectens (type species of Alilepus), A. laskarewi and A. hungaricus are designated here. The new material from Egorovka 2 (MN12) allowed a detailed morphometric re-description and emended diagnosis of A. laskarewi, one of the least understood species of the Old World Alilepus. One of the peculiarities of A. laskarewi is the very high incidence of mesofossettid in p3. The oldest leporids of the study area are verified for the MN11 localities Palievo, UA and Keinar, MD, but the taxonomy for these very scanty records remains unclear. In the light of the new material presented here, (i) a synoptic survey and discussion on the presence of p3 mesofossettid and (ii) hypotheses on a possible evolution and dispersal among the Old and New World Leporinae are provided.


ANGELONE, C. & ROOK, L. 2011. Alilepus meini n. sp., a new leporid (Leporidae, Lagomrpha) from the early Messinian of Tuscany (central western Italy). Geobios 44, 151-156.View article

AVERIANOV, A.O. 1996. The Neogene rabbit Hypolagus igromovi Gureev, 1964 (Lagomorpha, Leporidae) from southern European Russia. Acta zoologica cracoviensia 39(1), 61-66.

AVERIANOV, A.O. & TESAKOV, A.S. 1997. Evolutionary trends in Mio-Pliocene Leporinae, based on Trischizolagus (Mammalia, Lagomorpha). Paläontologische Zeitschrift 71, 145-153.View article

BERGGREN, W.A., KENT, D.V., SWISHER, C.C. & AUBRY, M.-P. 1995. A revised Cenozoic geochronology and chronostratigraphy, 129-212. In BERGGREN, W.A., KENT, D.V. & HARDENBOL, J. (eds) Geochronology Time Scales and Global Stratigraphic Correlations: A Unified Temporal Framework for a Historical Geology. SEPM Special Publication No. 54. SEPM, Tulsa.

BOON-KRISTKOIZ, E. & KRISTKOIZ, A.R. 1999. Order Lagomorpha, 259-262. In RÖSSNER, G. & HEISSIG, K. (eds) The Miocene Land Mammals of Europe. Verlag Dr. Friedrich Pfeil, München.

BRANDT, J.F. 1855. Beiträge zur näheren Kenntniss der Säugethiere Russlands. Mémoire de l’Académie impériale des Sciences, St. Petersburg, Physique, Mathématique, et Naturalistique Séries 6(9), 1-365.

BRUIJN, H. DE 1995. The vertebrate locality Maramena (Macedonia, Greece) at the Turolian-Ruscinian boundary (Neogene). 11 - Lagomorpha (Mammalia). Münchner Geowissenschaftliche Abhandlungen A 28, 133-136.

ČERMÁK, S. 2009. The Plio-Pleistocene record of Hypolagus (Lagomorpha, Leporidae) from the Czech and Slovak Republics with comments on systematics and classification of the genus. Bulletin of Geosciences 84(3), 497-524.View article

ČERMÁK, S. & WAGNER, J. 2013. The Pliocene record of Trischizolagus and Pliopentalagus (Leporidae, Lagomorpha, Mammalia) in Central Europe with comments on taxonomy and evolutionary history of Leporinae. Neues Jahrbuch fur Geologie und Paläontologie Abhandlungen 268, 97-111.View article

CHENG, J., CAO, B., TIAN, M. & LI, L. 1995a. A new early Pleistocene Mammalian fauna from Zhoukoudian. Earth Science (Wuhan) 20, 497-504.

CHENG, J., CAO, B., TIAN, M., YUAN, L., ZHAO, Z., LI, L., LI, C. & CHEN, L. 1995b. Early Pleistocene cave deposits with fossil mammals at Zhoukoudian. Geoscience 9, 441-449.

DAWSON, M. 1958. Late Tertiary Leporidae of North America. University of Kansas Paleontological Contributions 6, 1-75.

DAXNER, G. & FEJFAR, O. 1967. Über die Gattung Alilepus Dice, 1931 und Pliopentalagus Gureev, 1964 (Lagomorpha, Mammalia). Annalen des Naturhistorischen Museums Wien 71, 37-55.

DELINSCHI, A. 2014. Late Miocene lagomorphs from the Republic of Moldova. Annales de Paléontologie 100, 157-163.View article

DENG, T. 2006. Chinese Neogene mammal biochronology. Vertebrata Palasiatica 44, 143-163.

DIETRICH, W.O. 1941. Die säugetierepaläontologischen Erhebnisse der Kohl-Larsen’schen Expedition 1937-1939 in nördlichen Deutsch-Ostafrika. Zeitblatt Mineralogie B 8, 217-223.

DICE, L.R. 1917. Systematic position of several tertiary lagomorphs. University of California Publications, Bulletin of the Department of Geology 10, 179-183.

DICE, L.R. 1929. The phylogeny of the Leporidae, with description of a new genus. Journal of Mammalogy 10, 340-344.View article

DICE, L.R. 1931. Alilepus, a new name to replace Allolagus Dice, preoccupied, and notes on several species of fossil hares. Journal of Mammalogy 12, 159-160.View article

DOWNEY, J.S. 1968. Late Pliocene Lagomorphs of the San Pedro Valley, Arizona. United States Geological Survey, Professional Paper 600D, 169-173.

ERBAEVA, M.A. & ANGERMANN, R. 1983. Das Originalmaterial von Serengetilagus praecapensis Dietrich, 1941 - ergänzende Beschreibung und vergleichende Diskussion. Schriftenreihe für geologische Wissenschaften 19/20, 39-60.

FAHLBUSCH, V., QIU, Z.-D. & STORCH, G. 1983. The Neogene mammalian faunas of Ertemte and Harr Obo in Nei Mongol, China. - I. Report on field work in 1980 and preliminary results. Scientia Sinica B 26, 205-224.

FEJFAR, O. 1961. Die plio-pleistozänen Wirbeltierfaunen von Hajnáčka und Ivanovce (Slowakei), ČSSR. III. Lagomorpha. Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 1961(5), 267-282.

FEJFAR, O. & HEINRICH, W.-D. 1983. Arvicoliden-Sukzession und Biostratigraphie des Oberpliozäns und Quartärs in Europa. Schriftenreihe für Geologische Wissenschaften 19/20, 61-109.

FEJFAR, O., HEINRICH, W.-D. & LINDSAY, E.H. 1998. Updating the Neogene Rodent biochronology in Europe. Mededelingen Nederlands Instituut voor Toegepaste Geowetenschappen TNO 60, 533-554.

FEJFAR, O., HEINRICH, W.-D., KORDOS, L. & MAUL, L.C. 2011. Microtoid cricetids and the early history of arvicolids (Mammalia, Rodentia). Palaeontologia Electronica 14(3), 27A: 1-38;

FISCHER, J.G. 1817. Adversaria zoologica. Mémoires de la Société Impérialle Naturelle (Moscow) 5, 368-428.

FLADERER, F.A. 1987. Beitrag zur Entwicklung von Hypolagus und Lepus (Lagomorpha, Mammalia) im Pliopleistozän von Mitteleuropa. Sitzungsberichte der Österreichische Akademie der Wissenschaften, Mathematisch-naturwissenschaftliche Klasse, Abteilung I 196, 123-138.

FLADERER, F.A. & REINER, G. 1996. Evolutionary shifts in the first premolar pattern of Hypolagus beremendensis (Petényi, 1964) (Lagomorpha, Mammalia) in the Plio-Pleistocene of Central Europe. Acta zoologica cracoviensia 39, 147-160.

FLYNN, L.J., WINKLER, A.J., ERBAEVA, M., ALEXEEVA, N., ANDERS, U., ANGELONE, C., ČERMÁK, S., FLADERER, F.A., KRAATZ, B., RUEDAS, L.A., RUF, I., TOMIDA, Y., VEITSCHEGGER, V. & ZHANG, Z. 2014. The Leporid Datum: A late Miocene biotic marker. Mammal Review 44(3-4), 164-176.View article

FLYNN, L.J., WU, W.Y. & DOWNS, W.R. 1997. Dating vertebrate microfaunas in the late Neogene record of Northern China. Palaeogeography, Palaeoclimatology, Palaeoecology 133, 227-242.View article

GAZIN, C.L. 1930. A Tertiary vertebrate fauna from the upper Cuyama drainage basin, California. Carnegie Institution of Washington 404, 55-76.

GAZIN, C.L. 1934. Fossil Hares from the Late Pliocene of Southern Idaho. Proceedings of the United States National Museum 83(2976), 111-121.View article

GUREEV, A.A. 1964. Fauna of the USSR (Lagomorpha), Vol. 3 (10). 276 pp. Nauka, Moscow & Leningrad. [in Russian]

HIBBARD, C.W. 1939a. Four new rabbits from the upper Pliocene of Kansas. American Midland Naturalist 21(2), 506-513.View article

HIBBARD, C.W. 1939b. Nekrolagus, new name for Pediolagus Hibbard, not Marelli. American Midland Naturalist 21(3), Table of Contents.

HIBBARD, C.W. 1963. The origin of the p3 pattern of Sylvilagus, Caprolagus, Oryctolagus and Lepus. Journal of Mammalogy 44, 1-15.View article

HIBBARD, C.W. 1969. The rabbits (Hypolagus and Pratilepus) from the Upper Pliocene, Hagerman Local Fauna of Idaho. Papers of the Michigan Academy of Science, Arts, and Letters 1(1), 81-97.

HULVA, P., HORÁČEK, I. & BENDA, P. 2007. Molecules, morphometrics and new fossils provide an integrated view of the evolutionary history of Rhinopomatidae (Mammalia: Chiroptera). BMC Evolutionary Biology 7, 165, 1-28.View article

ICZN 1999. International Code of Zoological Nomenclature, 4th Edition. XXIX + 306 pp. International Trust for Zoological Nomenclature, London.

JIN, C.Z. 2004. Fossil leporids (Mammalia, Lagomorpha) from Huainan, Anhui, China. Vertebrata PalAsiatica 42, 230-245. [in Chinese, English summary]

JIN, C.Z., TOMIDA, Y., WANG, Y. & ZHANG, Y. 2010. First discovery of fossil Nesolagus (Leporidae, Lagomorpha) from Southeast Asia. Science China Earth Sciences 53, 1134-1140.View article

KHOMENKO, J. 1914. La faune méotique du village Taraklia du district de Bendery. Travaux de la Société des Naturalistes et des Amateurs des Sciences Naturelles de Bessarabie 5, 1-55.

KORMOS, T. 1911. Der pliozäne Knochenfund bei Polgárdi (Vorläufiger Bericht). Földtani Közlöny 41, 171-189.

KORMOS, T. 1934. Az euráziai nyulak származástani problémája - Zur Frage der Abstammung eurasiatischer Hasen. Allatani Közlemények 31, 65-78.

KOROTKEVICH, E.L. 1988. A history of foundation of hipparion fauna of Eastern Europe. 164 pp. Naukova dumka, Kiev. [in Russian]

KRETZOI, M. 1954. Befejező jelentés a Csakvári barlang őslénytani feltásáról [Final report on the paleontological findings in the cave of Csákvár]. A Magyar Állami Földtani Intézet Évi Jelentése az 1952 Évről 1952, 37-69. [in Hungarian]

LÓPEZ MARTÍNEZ, N. 1976. Lagomorpha from the Turolian of Pikermi (Greece). Proceedings Koninklijke Nederlandse Akademie van Wetenschappen 79, 235-244.

LÓPEZ MARTÍNEZ, N. 1977. Nuevos Lagomorfos (Mammalia) del Neógeno y Cuaternario espanol. Trabajos sobre Neógeno-Cuaternario 8, 7-45.

LÓPEZ MARTÍNEZ, N. 1989. Revisión sistemática y biostratigráfica de los Lagomorpha (Mammalia) del Terciario y Cuaternario de Espana. Memorias del Museo Paleontológico de la Universidad de Zaragoza 3, 1-342.

LÓPEZ MARTÍNEZ, N. 2008. The lagomorph fossil record and the origin of the European rabbit, 27-46. In ALVES, P.C., FERRAND, N. & HACKLÄNDER, K. (eds) Lagomorph Biology: Evolution, Ecology and Conservation. Springer-Verlag, Berlin & Heidelberg.

LÓPEZ MARTÍNEZ, N., LIKIUS, A., MACKAYE, H.T., VIGNAUD, P. & BRUNET, M. 2007. A new Lagomorph from the Late Miocene of Chad (Central Africa). Revista Espanola de Paleontología 22, 1-20.

LUNGU, A. 1980. New records of the Sarmatian land vertebrate fauna in Moldavia, 3-8. In Quaternary and neogene faunas and floras of Moldavian SSR. Shtiintsa, Kishinev. [in Russian]

LUNGU, A. & RZEBIK-KOWALSKA, B. 2011. Faunal assemblages, stratigraphy and taphonomy of the Late Miocene localities in the Republic of Moldova. 62 pp. Institute of Systematics and Evolution of Animals Polish Academy of Sciences, Krakow.

LUNGU, A., DELINSCHI, A. & NICOARA, I. 2007. Some dates about studies representative order Lagomorpha from hipparion fauna, of superior Miocen from Republic of Moldova. Scientific Bulletin of National Museum of Ethnography and Natural History of Moldova. Ethnography, Natural Sciences and Museology 6(19), 141-143.

MEIN, P. 1988. Continental faunas at the Miocene-Pliocene boundary. La gisement karstique turolien de Lissieu (Rhône). International Workshop “Continental Faunas at the Miocene/Pliocene Boundary”. Proceedings, Faenza, 25.

MEIN, P. 1999. The late Miocene small mammal succession from France, with emphasis on the Rhône Valley localities, 140-164. In AGUSTÍ, J., ROOK, L. & ANDREWS, P. (eds) The Evolution of Terrestrial Ecosystems in Europe, Volume 1. Hominoid Evolution and Climatic Change in Europe. Cambridge University Press, Cambridge.

MILLER, W.E. & CARRANZA-CASTANEDA, O. 1982. New lagomorphs from the Pliocene of central Mexico. Journal of Vertebrate Paleontology 2, 95-107.View article

NESIN, V.A. 1996. Lower Pliocene rodents of Ukraine and problems of Pontian biostratigraphy. Acta zoologica cracoviensia 39(1), 395-399.

NESIN, V.A. 2004. Stratigraphical position of the Late Neogene localities of the microtheriofauna in Ukraine, 195-199. In GOZHI, P.F. (ed.) Problemy stratigrafii i fanerozoyu Ukrainy. Institut geologichnikh nauk, Kiev. [in Russian]

NESIN, V.A. 2013. The Neogene Murinae (Rodentia, Muridae) of Ukraine. 174 pp. University book, Sumy. [in Russian, English summary]

NESIN, V.A. & NADACHOWSKI, A. 2001. Late Miocene and Pliocene small mammal faunas (Insectivora, Lagomorpha, Rodentia) of Southeastern Europe. Acta zoologica cracoviensia 44(2), 107-135.

OGNEV, S.I. 1929. Zur Systematic der russischen Hasen. Zoologischer Anzeiger 84, 68-83.

PALACIOS, F. & LÓPEZ MARTÍNEZ, N. 1980. Morfología dentaria de las liebres europeas (Lagomorpha, Leporinae). Donana Acta Vertebrata 7, 61-81.

QI, G.-Q., DONG, W., ZHENG, L., ZHAO, L., GAO, F., YUE, L.-P. & ZHANG, Y.-X. 2006. Taxonomy, age and environment status of the Yuanmou hominoids. Chinese Science Bulletin 51, 704-712.View article

QIU, Z.-D. 1987. The Neogene mammalian faunas of Ertemte and Harr Obo in Inner Mongolia (Nei Mongol), China. 6. Hares and pikas - Lagomorpha: Leporidae and Ochotonidae. Senckenbergiana lethaea 67, 375-399.

QIU, Z. & HAN, D. 1986. Fossil Lagormorpha from the hominoid locality of Lufeng, Yunnan. Acta Anthropologica Sinica 5, 41-53. [in Chinese, English summary]

QIU, Z. & QIU, Z. 1995. Chronological sequence and subdivision of Chinese Neogene mammalian faunas. Palaeogeography, Palaeoclimatology, Palaeoecology 116, 41-70.View article

QIU, Z., WU, W. & QIU, Z. 1999. Miocene mammal faunal sequence of China: paleozoogeography and Eurasian relationships, 443-445. In RÖSSNER, G.E. & HEISSIG, K. (eds) The Miocene Land Mammals of Europe. Verlag Dr. Friedrich Pfeil, München.

RABEDER, G. 1989. Die Säugetiere des Pontien in Österreich und Ungarn. Jugoslawische Akademie der Wissenschaften und Künste 8, 821-836.

RADDE, G. 1861. Neue Säugethier-Arten aus Ost-Sibirien. Mélanges Biologiques tirés du Bulletin Physico-Mathematique de l’Académie Impériale des Sciences de St. Petersbourg 3, 676-687.

RADULESCO, C. & SAMSON, P. 1967. Contribution a la connaissance du complexe faunistique de Malusteni-Beresti (Pléistocene inférieur), Roumanie I. Ord. Lagomorpha, Fam. Leporidae. Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 1967(9), 544-563.

RUSSELS, B.D. & HARRIS, A.H. 1986. A new leporine (Lagomorpha: Leporidae) from Wisconsinan deposits of the Chihuahuan Desert. Journal of Mammalogy 67, 632-639.View article

SCHLOSSER, M. 1924. Tertiary vertebrates from Mongolia. Palaeontologia Sinica, Series C 1(1), 1-133.

ŞEN, Ş. 1983. Rongeurs et lagomorphes du gisement Pliocene de Pul-e Charkhi, basin de Kabul, Afghanistan. Bulletin du Museum National d’Histoire Naturelle, Paris, 5e séries, 5C(1 ), 33-74.

ŞEN, Ş. & ERBAEVA, M. 1995. Early Pliocene leporids (Mammalia, Lagomorpha) from Afghanistan. Comptes Rendus de l’Académie des Sciences de Paris, Sér. II 320, 1225-1231.

SEMENENKO, V.N. 1987. [Stratigraphical Correlation of the Upper Miocene and Pliocene of the Eastern Paratethys and Tethys]. 230 pp. Naukova dumka, Kiev. [in Russian]

SINITSA, M.V. 2005. The small mammal fauna (Insectivora, Lagomorpha, Rodentia) from the Meotian deposits of Kuyalnik River. Proceedings of the museum Brauner’s foundation 11(4), 11-17. [in Russian]

SINITSA, M.V. 2008. Meotian small mammals from the Egorovka locality, 285-289. In GOZHIK, P.F. (ed.) Biostratigraphic fundamentals of creating the stratigraphic schemes of the Phanerozoic of Ukraine. Proceedings of the Institute of Geological Sciences of the NAS of Ukraine, Kyiv. [in Russian, English summary]

SINITSA, M.V. 2009a. Cricetids (Rodentia, Mammalia) from the Upper Miocene of Egorovka locality. Vestnik zoologii 44, 209-225. [in Russian, English summary]

SINITSA, M.V. 2009b. A new small mammal fauna from the Lower Turolian (MN11) of the Southern Ukraine, 181-182. In CODREANU, I. (ed.) Materialele simpozionului jubiliar internaţional ‘Mediul şi dezvoltarea durabilă’. Labirint, Chişinău.

SINITSA, M.V. 2011. Pliopetaurista (Mammalia, Rodentia) from the Neogene of Ukraine. Vestnik zoologii 45, 19-33. [in Russian, English summary]

SINITSA, M.V. 2012. Cricetids (Mammalia, Rodentia) from the Late Miocene locality Palievo, Southern Ukraine. Vestnik zoologii 46, 137-147. [in Russian, English summary]

STEININGER, F.F. 1999. The Continental European Miocene. Chronostratigraphy, Geochronology and Biostratigraphy of the Miocene “European Land Mammal Mega-Zones” (ELMMZ) and the Miocene “Mammal-Zones (MN-Zones)”, 9-24. In RÖSSNER, G.E. & HEISSIG, K. (eds) The Miocene Land Mammals of Europe. Verlag Dr. Friedrich Pfeil, München.

SYCH, L. 1965. Fossil Leporidae from the Pliocene and the Pleistocene of Poland. Acta zoologica cracoviensia 10, 1-88.

TOMIDA, Y. 2008. Can morphological differences among extinct species be explained by means of genetics? An example from fossil rabbits. Journal of Vertebrate Paleontology 28 (3, Supplement), 152A.

TOMIDA, Y. & JIN, C.-Z. 2009. Two new species of Pliopentalagus (Leporidae, Lagomorpha) from the Pliocene of Anhui Province, China, with a revision of Pl. huainanensis. Vertebrata PalAsiatica 47(1), 53-71.

TOPACHEVSKI, V.A. & SKORIK, A.F. 1992. The Neogene and Pleistocene lower cricetids of SE Europe. 242 pp. Naukova dumka, Kiev. [in Russian, English summary]

TOPACHEVSKI, V.A., NESIN, V.A. & TOPACHEVSKI, I.V. 2000. A small mammal oryctocenoses in the Meotian of the Cherevichnoe section. Proceedings of the National Academy of Sciences of Ukraine 10, 192-195. [in Russian, English summary]

VOORHIES, M.R. & TIMPERLEY, C.L. 1997. A new Pronotolagus (Lagomorpha: Leporidae) and other leporids from the Valentine Railway Quarries (Barstovian, Nebraska), and the archaeolagine-leporine transition. Journal of Vertebrate Paleontology 17, 725-737.View article

WHITE, J.A. 1991. North American Leporinae (Mammalia: Lagomorpha) from late Miocene (Clarendonian) to latest Pliocene (Blancan). Journal of Vertebrate Paleontology 11, 67-89.View article

WHITE, J.A. & MORGAN, N.H. 1995. The Leporidae (Mammalia: Lagomorpha) from the Blancan (Pliocene) Taunton Local Fauna of Washington. Journal of Vertebrate Paleontology 15, 366-374.View article

WILSON, R.W. 1938. A new genus of lagomorph from the Pliocene of Mexico. Bulletin of the Southern California Academy of Sciences 36(3), 98-104.

WINKLER, A.J. & AVERY, M. 2010. Lagomorpha, 305-317. In WERDELIN, L. & SANDERS, W.J. (eds) Cenozoic Mammals of Africa. University of California Press, Berkeley.View article

WINKLER, A.J. & TOMIDA, Y. 2012. The fossil record of African leporids. 4th World Lagomorph Conference, Proceedings, Vienna, 139.

WINKLER, A.J., FLYNN, L.J. & TOMIDA, Y. 2011. Fossil lagomorphs from the Potwar Plateau, northern Pakistan. Palaeontologia Electronica 14(3), 38A, 1-16;