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Part 1 Gennady+G. Boeskorov of the of Pliocene and Pleistocene A review systematics moose, part 1 A review of the systematics of Pliocene and Pleistocene moose, part 1 Gennady+G. Boeskorov Summary of evolutionand of the from The results investigations on the the systematics moose are presented. New data our own research and data from the literature on morphological variability and genetics of the living moose are discussed. In data fossil broad-fronted Cervalces and is submitted. The former addition, new on moose ( ) true moose (Alces) genus and be direct of Alces. Based the data revision of the of the Libralces appear not to ancestors upon new a systematics moose is carried out, and questions on philogeny are considered. Samenvatting onderzoek de De resultaten van naar evolutie en de systematiek van de eland wordenin ditartikel gepresenteerd. uit variatie Gegevens afkomstig van eigen onderzoek en de literatuur over morfologische en erfelijkheid van de eland worden Daarnaast worden nieuwe de fossiele hert-eland Cervalces huidige besproken. gegevens van en van fossiele elandAlces Cervalces Libralces directe voorouders Alces. de ware gepresenteerd. en blijken geen te zijn van Aan hand nieuwe wordt revisie ondernomen de de worden van gegevens een van systematiek van eland, en vragen betreffende hun stamboom behandeld. Introduction The systematic position of the modern moose is under discussion. Within their huge Holarctic habitat, the moose is represented by a number The of the terrestrial animals moose, one largest of forms with various of morphological of the Northern and degrees Hemisphere an important isolation. The prevails that Northern has opinion hunting object, a large economic importance Eurasia and North America is occupied by one in Sweden, Finland, in Norway, many regions species - Alces alces L. which contains at most of Russia, the USA and Canada. The original eight subspecies: the moose or elk (A. of Alces alces European English name is 'elk', but nowa- a. alces), the East-Siberian moose (A. a. pfitzen- the American is days name, 'moose', usually mayeri), the Manchurian or Ussuri moose (A. a. used in scientific literature. The moose is the cameloides), Buturlin's moose (A. a. buturlini), the largest representative of the family of deer. It , Alaskan moose (A. a. gigas), the West-Canadian essentially differs from the other deer by its moose a. andersoni), the East-Canadian habitus: but with rather (A. very massive a short and the Yellowstone moose (A. a. americanus), trunk, a strongly developed thorax, a large, moose (A. a. shirasi) (Peterson, 1955,1974; and elongated hump-nosed head, a very large Sokolov, 1959; Heptner et al. 1961; Wilson & and massive and wide r upper lip, very large Reeder, 1992; Geist, Thefirst and which 1998). description ears, very long legs, a long 'bell', is a of the Cervus moose as a species on its own ( hanging outgrowth of skin under the throat. alces) was made by Karl Linne in 1758, based on The antlers in the male project laterally from the the moose from Sweden; gradually this name head and are usually palmated (shovel-shaped became used for other moose populations as expanded). The front part of the moose torso is well. In 1821 Gray attributed the species to a more massive than the hind part. This characte- Alces. Since the description of ristic also of separate genus: constitution causes a peculiar way the American moose as Cervus americanus movement, with a quick, round trot, a pliable Clinton considered it 1822, some a separate step, at which the trace of the hind limbs blocks species, distinct from the European A. alces a trace of the front limbs. The lengthened limb (Miller, 1899; Seton, 1910; Lydekker, 1915; in the bones, particular metapodials, give it a Anthony, 1928). An interesting detail is that high step, which promotes movement on tracks during the 19th century several times moose that are almost impassable for other deer: were imported to England from Canada; some hillocky bogs, swamps, wind-fallenwood and experts (e.g. Rolfe, 1983) even identified them deep snow. with the extinct giant deer, or Irish elk, Megalo- 26 CRANIUM 22, 2 - 2005 ceros giganteus. Unfortunately Linne himself Yakut Institute of the Geological Sciences Sibe- never saw the American moose. rian Division of the RAS at Yakutsk (YIGS), Mammoth Museum of Yakutsk (MM), GM, Based features of the upon some morphological PMM, American Museumof Natural History of Flerov indicated some diffe- moose, (1931,1934) New York (AMNH), IRMN, VMN, museums of rences between moose from Europe and North natural history in Betenkes (MB) and Chersky America, and concluded that East Siberian (CMN) and in the Museum of East-Yakutian moose are closer to the latter than to European United Mine-Geological organization at Khan- moose. He proposed two separate moose dyga (MEYUMGO). Some fossil and subfossil species: A. alces L., inhabiting Scandinavia, part remains of Alces were found by us during work of Eastern Europe, and Siberia westward of the on the Rebrovo outcrop (the Laptev strait sea- Yenissey river, andA. americanus Clinton, inha- shore, 1995) and Duvanny Yar outcrop (lower biting North America, Siberia eastward of the stream of the Kolyma river, 1995,1997,1999). Yenissey, and the Russian Far East. Although Of fossil and subfossil Alces we studied 19 some scholars agreed with this (e.g. complete skulls, 14 lower jaws, 25 antler frag- Vereshchagin, 1949), Flerov later stated that the ments, and 4 metatarsals; of Cervalces we observed differences did not exceed subspecies studied 13 partial skulls, 8 lower jaws, 31 antler level (Flerov, 1952). Nevertheless, his earlier beams and 6 metatarsals. had influence and Russian opinion large many mammalogists considered the moose from East Skull measurementswere made following and Northeast Siberia to belong to the American Sokolov (1959) with some additions. We measured the skull in moose (Bobrinsky et al., 1944; Gromov et al., parameters as given 1963; Baryshnikov et al., 1981). The consensus at figure 1. The lower jaw parameters are: 1. total front of the without the moment is that the American moose differs length (from the edge jaw, level back of mandibular 2. from European mooses at species (Gromov teeth, up to edge angle; et al., 1963; Chernyavsky & Domnich, 1989) or coronar length of the tooth row; 3. alveolar of of semi-species level (Groves & Grubb, 1982). length the tooth row; 4. length premolar 5. of molar 6. of Subsequently, differences in chromosome row; length row; length numbers, and in a number of other genetic, diastema; 7. height from the lower edge of the mandibular the 8. ecological and ethological features were found angle to coronar process; between the European and North American height of the mandibleahead P2; 9. height of the of antlers moose. Although morphologic and genetic mandible behind M3. Measurements differences between the European and (spread, length of antler, width of palmation) American moose were discussed several times was carried out according to the rules of the (Filonov, 1983; Bubenik, 1986; Chernyavsky & International Union of the Hunters. Only antler Domnich, 1989; Danilkin, 1999), the question on tines longer than 5 cm were taken into account. the number of species and subspecies of Length of antler beam (shaft) was measured modern moose remained unsolved till now. from the burr prior to the beginning of palma- tion, and beam circumference was measured In order to solve this problem we started above the burr and at midpoint (fig. 2). research on the systematics, caryology and morphological variability of the moose in 1992. this article the In we present our analysis on of differentiation of the European, Siberian, the Evolutionary history moose Russian Far Eastern and American moose; part Fossil and modern moose are attribted to a of our results has already been published (Boes- separate subfamily: Alcinae Jerdon, 1874 (Frick, korov, 1996,1996a, 1997,1998; Boyeskorov, 1937; Haltenorth, 1963; Sher, 1986; Vislobokova, 1999). To solve some controversial points in the 1990). The origin of this subfamily lies some- of the we included material philogeny moose, where in the Early Pliocene (5,5 - 4,1 million Alces and of the extinct of fossil and subfossil this the Pseudalces years ago). To period original broad-fronted moose Cervalces. mirandus Flerov, 1962 belongs, which was described from the Ciscaucasia (Kosyakinsky This ancient quarry, near Stavropol). 'pseudo- Material and Methods -moose' had a skull with an elevation between the basements of the antlers. Its antler beams Fossil and subfossil remains of Alces and fossil were directed laterally, backwards and slightly broad-fronted moose Cervalces were studied in: 27 A review of the of Pleistocene 1 systematics Pliocene and moose, part of - - - 1 Measurements skull. 1 2 condilobasal 3 cheek tooth 4 - widthof Fig a moose length; length; upper row; cheek bones; 5 - rostrum length (from front edge of intermaxillar bones to connecting point of nasal bones and - maxillarbones); 6 orbito-facial length; 7 - length of nasal bones; 8 - skull height in the nasion point (from the posterior edge of the nasal bones perpendicularly to the skull axis to the lower edge of the maxillar bone, without - 9 rostrum 10 - from the lower of foramen 11 - from teeth); width; occipital height edge magnum; occipital height the of 12 - maximum 13 - width of upper edge foramenmagnum; occipital width; occipital condyles Metingen aan de elandschedel. 1 - lengte; 2 - condilobasalelengte; 3 - rij bovenkaakskiezen; 4 - breedte ter hoogte de 5 - de intermaxillarebeenderen het van jukbeenderen; snuitlengte (gemeten van voorste punt van tot con- de - - tactpunt van neusbeenderenmet dekaakbeenderen); 6 orbito-faciale lengte; 7 lengte van de neusbeenderen; 8 - in het de achterrand de neusbeenderenloodrecht schedelhoogte nasion-punt (gemeten van van op de sche- delas de onderrand het zonder 9 - - tot van kaakbeen, gebitselementen); snuitbreedte; 10 achterhoofdshoogte van- af de onderrand - van het achterhoofdsgat; 11 achterhoofdshoogte vanaf de bovenrand van het achterhoofdsgat; - 12 maximale achterhoofdsbreedte; 13 - breedte van de achterhoofdsknobbels 28 CRANIUM 22, 2 - 2005 - - of antlers.
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