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Part 2 Gennady+G. Boeskorov CRANIUM 23,1 - 2006 A review of the systematics of Pliocene and Pleistocene moose, part 2 Gennady+G. Boeskorov Summary This is the second ofthe article theevolution and the ofthe Data from the literature paper part on systematics moose. and of the and data from research New onmorphological variability genetics moose new our own are discussed. data and on fossil broad-fronted moose (Cervalces; see part 1) true moose (Alces) is submitted. The former and Libralces direct of.Alces In of revision ofthe of the carried (part 1) are no ancestors (part 2). light new data, a systematics moose is out, and questions on phylogeny are considered. Samenvatting Dit artikel is het tweede deel de de evolutie de de eland. van publicatie over en systematiek van Gegevens uit de variatie de eland onderzoek worden literatuur over morfologische en erfelijkheid van en gegevens van eigen Ook worden de hert-eland Cervalces fossiele eland besproken. nieuwe gegevens van fossiele (deel 1) en van ware Alces Cervalces Libralces Aan (dit deel) gepresenteerd. en blijken geendirecte voorouders te zijn van Alces. de handvan nieuwe wordt revisieondernomen de de eland worden betreffendehun gegevens een van systematiek van en vragen phylogenie behandeld. Family Cervidae Gray,1821 sediments of theOb river (Krasny Yar) are attri- buted to true moose (Alexeeva, 1980). Moose bones found in morrain on the Kola peninsula Subfamily Alcinae Jerdon, 1874 probably belong to the Middle Pleistocene Genus Alces Gray, 1821 (Gromova, 1965; Vereshagin, 1967). To the late Cervus alces 1758. Type species. Linnaeus, Middle Pleistocene (Tasovskoe glaciation) true Present, in Europe. moose are assigned connection with a fauna at the Lower Antler beam of periglacial (tundra-steppe) Diagnosis. average length or Tunguska river (Vangengeim, 1977). short (length always less than circumference). Braincase is rather narrow and than in higher Pavlov (1906) described an adult true moose Cervalces. The facial of the skull portion is long, skull foundat the Kama river in the vicinity of and the rostrum is Nasal strongly elongated. Mysy village under the name “Alces savinus bones their less than that of are short; length is Fisch." This skull to (fig. 19p). may belong a the dental The of row. upper (nasal) processes Khasarian fauna of the Middle Pleistocene the bones do reach the premaxillary usually not (Gromova, 1965), though its stratigraphic posi- nasal bones. The between the antlers bears area tion requires a re-examination (Sher, 1986). The a cone-like The orbits pronounced prominence. skull shows characteristics of Alces: a long almost level with the frontal vault. are on a rostrum and short nasal bones. Its upper of bones processes premaxillary are widened, as Species composition. At least two species are in modern European moose. The antler beam is recognised: A. alces (Linnaeus, 1758), early Late long, though much shorter than in broad- Pleistocene - present, Western Eurasia; A. ameri- fronted antlers close the moose; are to deer-like canus (Clinton, 1822), early Late Pleistocene - type with only a few tines. An original feature present, Eastern (?) Eurasia, end of the Late of this moose is a well-expressed tine on the Pleistocene - North (Wisconsinan) present, right beam (fig. 19p). The development of an America. additional tine on the beam is not characteristic Alces sp. for modern moose: a similar formation is not in found modernAlces, nor of moose by Vereshchagin Remains true (Alces) are found in 136 pairs of antlers), nor us deposits from the late Middle Pleistocene (1949; among by more than 300 of onwards. Finds of Alces in the Netherlands (among pairs antlers). Appa- sp. rently, there was an original early Pleistocene (Erdbrink, 1954) belong to that period. Some Alces for which the of an additional moose remains from the MiddlePleistocene presence tine on the beam was characteristic. Similar 3 review of the of Pliocene and Pleistocene 2 A systematics moose, part Fig 19 Antlers of the fossil and subfossil Alces from Western and Eastern Europe: a - Germany, Schussenried, Late Würm after b - - (Kahlke, 1956, Sher, 1971); specimen at PMM, Germany, Bavaria, Holocene; c AMNH 27772, d - - k - Denmark, Holocene; specimen at PMM, Austria, Brück, Holocene; e “Alces palmatus”, Romania, Pleistocene (after Macarovici, 1961); 1, m - Western Ukraine, Pleistocene (after Vereshchagin, 1967); n - of - North-East Russia, the Kichmenga river, Pleistocene (?) (after Vereshchagin & Russakov, 1979); o “Alces of Late Pleistocene-Holocene and - “Alces middlestream of fossilis”, near Moscow, vicinity Mytishchi, p savinus”, the Volga river, lateMiddle Pleistocene (?) (both - after Pavlow, 1906). Geweienvan fossiele en subfossiele Alces uit West- en Oost-Europa: a - Duitsland, Schussenried, Laat Würm (Kahlke, 1956,naar Sher, 1971); b - stuk uit PMM, Duitsland, Beieren, Holoceen; c - AMNH 27773, Denemarken, d - uit - - Pleistoceen Holoceen; stuk PMM, Oostenrijk, Brück, Holoceen; e k “Alces palmatus”, Roemenië, (naar Macarovici, 1961); 1, m - West Oekraïne, Pleistoceen (naar Vereshchagin, 1967); n - Noordoost Rusland, Kichmen- - “Alces ga rivier, Pleistoceen (?) (naar Vereshchagin & Russakov, 1979); o fossilis”, omgeving van Moskou, nabij Laat - “Alces de Mytishchi, Pleistoceen-Holoceen en p savinus”, middenstroom van Wolga, laat MiddenPleisto- ceen (?) (beiden naar Pavlow, 1906). 4 CRANIUM 23,1 - 2006 Antlers of the fossil - Fig 20 and subfossil Alces: a specimen at GM RAS, near Moscow, vicinity of Mytishchi, Ho- - locene; b, c - GM RAS, Eastern Europe, Holocene (?); d ZM MSU, vicinity of Moscow (?), Pleistocene-Holocene; e - Stavropol district, near Caucasus region, Holocene (after Vereshchagin, 1949); A. alces cf. caucasicus: f - lower stream of the Don Middle - Krasnodar the - af- river, Sarkel fortress, Ages, g district, Urup river, Holocene (both i - Western Pleistocene ter Vereshchagin, 1959); h, Alces sp., Siberia, Krasny Yar, (after Alexeeva, 1980). Geweien fossiele Alces: - stuk in GM van en subfossiele a RAS, omgeving Moskou, nabij Mytishchi, Holoceen; b, c - GM RAS, Oost-Europa, Holoceen (?); d - ZM MSU, nabij Moskou (?), Pleistoceen-Holoceen; e - Stavropol dis- de f - trict, nabij Kaukasus, Holoceen (naar Vereshchagin, 1949); A. alces cf. caucasicus: benedenstroom van de Sarkel - Don, kasteel, Middeleeuwen, g district Krasnodar, Urup rivier, Holoceen (beiden naar Vereshchagin, i - Alces West Pleistoceen 1959); h, sp., Siberië, Krasny Yar, (naar Alexeeva, 1980). 5 review of the of Pliocene and Pleistocene 2 A systematics moose, part beams of true found The antler moose were by process among moose. wide, bi-partite Alexeeva (1980) in the Pleistocene sediments of palmation and elongated antler beam is charac- Western Siberia (fig. 20h-i). Vereshchagin (1959; teristic for the East-Siberian moose of the Late 1967) described the skull of fossil or subfossil Pleistocene. Most probably, the American moose from Cis-Caucasia with rudimental brow moose evolved in Eastern Siberia. tines (fig. 20e). Pleistocene Alces had, on Alces cf. alces and A. alces L., 1758 - moose of average, longer antler beams than modern the European type and the European moose moose (Vereshchagin, 1967). Cervus (Alces) fossilis H. V. Meyer, 1832. Eurasia until True moose were widespread in Alces palmatus Gray, 1843. the of Late Pleistocene beginning (see above). Alces leptocephalus Push, 1846. Probably, during this period their differentia- Material and locality. Separate antlers and frag- tion into two forms, the European and the ments: PMM 1955, Western Germany, Baden- East-Siberian, began. The latter later gave rise to Wurtemberg, Markt-Schwaben; PMM 1950/92, difficult to the North American moose. It is say Bavaria,. Taufkirchen (fig. 19b); PMM 1974/253, where this took In Late exactly process place. Bavaria, Kiinzing; PMM specimen without Pleistocene deposits, true moose remains are number, Austria, Bruk, Markt Grating, (fig. from the found Western Europe up to Russian 19d); AM 27772, Denmark, AMNH (fig. 19c); Far East and Japan (after Kahlke, 1999). Many GM 1823 and GM 1824, Eastern Europe (?), GM finds testify that they were distributed more RAS (fig. 20a, c). Skull fragments with antler widely than at present (see fig. 6 of part 1). parts: ZM 1411, vicinity of Mytishchi Sity (fig. antler for ZM without of A high variation in shape was typical 20a); specimen number, vicinity Pleistocene Alces (especially in western regions). Moscow (?), ZMMSU (fig. 20d); GM 1829 (?), of the Nerekhta Four antler types are recognized: shovel-shaped vicinity Kostroma Sity, river antlers with undivided palmation, and GM specimen without number, Tiraspol shovel-shaped antlers with bi-partite palmation, (?)• deer-like antlers, and antlers with an additional Geological age. The beam of the left antler the beam. this undoub- tine on In our opinion, PMM 1974/253 is found in the Pleistocene tedly testifies the existence of a differentiation Table 14 Sizes of fossil and subfossil Alces antlers in of the (in mm); case multiple specimens, averages are given. * ** Notes: - after specimens from Paleontological Museum of Munich (Germany) and AMNH (USA); - after specimens from Zoological Museum MSU and Geological Museum RAS, Moscow. Maten het is in van fossiele en subfossiele elandgeweien (in mm); gemiddelde gegeven het geval van meerdere * geweien. Opmerkingen: - naar geweien in het Paleontologisch Museumvan München (Duitsland) en AMNH ** - in Museum MSU (USA); naar geweien het Zoologische en het Geologische Museum RAS, Moskou. A. cf. america- AlcesAlces
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