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. - 2 3 shovel Fig 2 Scheme of measurements moose 1 spread; maximal antler length; (palmation)

- of at width; 4 - beam (shaft) length; 5 - circumference of beam above the burr; 6 circumference beam midpoint; 7

- circumference of burr

- Schema 1 - 2 - maximale 3 - 4 stok- van metingen aan elandgeweien. spreiding; gewei-lengte; palmatie-breedte;

- - - stok boven de 6 omtrek de stok 7 (schacht-)lengte; 5 omtrek van de (schacht) rozenkrans; van halverwege; omtrek van de rozenkrans

it of Alcinae also shows the characteristic upwards (Vislobokova, 1990). However, genus observed the whole might be that Pseudalces was not the oldest features that are during of but of the deer of the subfamilia. These genus Alcinae, giant Megace- evolutionary lineage dead features rini andmost probably presents a ending constant are, amongst others, branch of the subfamilia Cervinae (Flerov & horizontally diverged antler beams without

and structure Shevyreva, 1963). Remains of Alcinae gen. tines ending in palmation, specific indet. were found in the Late Pliocene (3,4 to 2,5 of teeth and limb bones, and telemetacarpals. million Late Pliocene and Pleistocene remains of years ago; Early-Middle Villafranchian) Early deposits of Transbaikalia, Udunga (Vislobokova Libralces were also found in other sites of the et al., 1995). Western Europe (England, Germany). In

former USSR the oldest remains of Libralces The first well-defined of Alcinae is the genus were found in the Chaprov fauna from the broad-fronted moose Libralces. It was described the the Liventsov quarry in region near Asov from the Late Pliocene (Late Villafranchian) Sea, dated about 2,5 million years (Bajgusheva, fauna of Seneze (France) as Libralces gallicus 1971) and in the Middle Villafranchian (Azzaroli, 1952). This species had an unusual Kuruksay fauna from Tadjikistan (Vislobokova, habitus for moose: relatively small withers' 1988). height (about 1,4 m), and a very long (up to 1 m) but thin and S-shaped, curved antler beam. Vereshchagin (1957) described another repre-

The palmation of its antlers was minimal, the sentativeTamanalcesof ancient moose: cauca- skull low and wide with thick sicus from the Tamanian fauna of the latest was very bones, Pleistocene from the nasal bones were long and reached the Pliocene or Early Taman

but this taxon be premaxillary bones (fig. 3a; characteristic peninsula (Tsymbal), may invalid. feature for the representatives of the subfamilia considered

Cervinae), the Ml and sometimes the M2 had the Paleomeryx fold. Nevertheless, this oldest

29 of the of Pliocene and Pleistocene 1 A review systematics moose, part

Fig 3 The skull lateralview: a - Libralces gallicus, Sénèze; b - C. latifrons, Süssenborn; c - C. scotti, New Jersey,

d - Cervus - “A lces savinus” 1846 Mount-Hermon(after Sher, 1986); elaphus (after Sokolov, 1979); e (after Rouillier,

- and from in the PIN f - Alces A. alces caucasicus Naniev, h A. americanus a specimen RAS); alces; g - (after 1956);

- - De - b C. C. New schedel in zij-aanzicht: a Libralces gallicus, Sénèze; latifrons, Süssenborn; c scotti, Jersey,

Mount-Hermon(naar Sher, 1986); d - Cervus elaphus (naar Sokolov, 1979); e - “Alces savinus”(naar Rouillier, 1846

schedel in PIN f - Alces alce - A. alces caucasicus h - A. americanus en een RAS); s; g (naar Naniev, 1956);

From the Early Pleistocene of the Forest Bed body weight was about 1000 kg. The huge

(Norfolk, England) the giant broad-fronted antler spread of this moose points to an inhabi-

moose Alces was described tant of The latifrons (Johnson, open landscapes (fig. 4a). latifrons

1874). Numerous finds of this moose revailed its form is distinguished from the true moose Alces

close with Libralces nasalia that features: antler beam and similarity : long by some very long contact the premaxillary bones (fig. 3b), hori- nasalia, shape of nasalia, contact between nasal zontally projected long antler beams, etc. On the and premaxillary bones, large occipital width, other hand, A. latifrons differs considerably etc. These significant differences favours the from Libralces: the antler beam of the former is identification of latifrons as a representative of shorter but of (up to 0,6 m) sufficiently more the genus Cervalces, not Alces (Azzaroli, 1981; massive, the facial length of the skull is longer, Sher, 1986; Vislobokova, 1990; Boeskorov, 2001).

and the former lacks canines and the upper However, there is no accordance on the generic Paleomeryx-fo ld in its molars. The giant broad- systematics of the subfamilia Alcinae. Vislo- has skull and fronted moose a very large bokova with Azzaroli that (1990) agrees (1952) postcranial skeleton: it is the largest form Libralces and considers three is a separate genus, fossil and living Cervidae and its among Alcinae: genera within Libralces, Cervalces and its withers' height probably exceeded 2,5 m and Alces. But later Azzaroli himself (Azzaroli, 1981;

30 CRANIUM 22, 2 - 2005

bokova, 1986,1996). Remains of C. latifrons and

C. cf. latifrons were found in Early Pleistocene deposits of Transbaikalia in the faunas of Kloch

nevo (Kozhamkulova, 1974; Vangengeim, 1977; Sher, 1986) and Zasukhino (Vangengeim &

Sotnikova, 1981). In the North-East of Siberia, the Kolyma lowland, the oldest remains of

broad-fronted moose were found in the Olye-

fauna - million rian (1,2 0,5 years ago) (Sher,

1971,1986). C. latifrons remains were also found

in the sediments at Adycha, which is correlated

to the Late Pliocene-Early Pleistocene (Anony-

Pleistocene - Middle mous, 1987). Early early

Pleistocene remains of C. cf. latifrons were also

found in North America in the Old Crow fauna,

analogous to the Olyerian fauna (Harington, 1978 in Sher, 1992).

In the Quaternary sediments of Northern

Eurasia (mainly in Eastern Siberia) antler frag-

ments are found with the original proportions: comparatively long (20 to 35 cm) but slender

14 to 17 antler beams. An - (circumference cm) Fig 4 Shape and sizes of Cervalces antlers: a C. lati- of the concomitant condi- frons (after Bubenik, 1986); b - C. postremus (resto- analysis geological the red); c - C. scotti (after Bubenik, 1986) tions and fauna reveals that at least majority of these finds belong to the late Middle Pleisto-

Cervalces - C. la- Vorm en van geweien: a afmetingen cene (Russanov, 1968; Sher, 1986). The new

tifrons (naar Bubenik, 1986); b - C. postremus (her- subspecific name Alces latifrons postremus was c - C. scotti Bubenik, 1986) steld); (naar for this with suggested moose relatively long but gracile antler beams (Vangengeim & Flerov,

1965). Others (Sher, 1986; Bubenik, 1986; Boes- followed Sher, 1986) included all broad- by korov, 2001) consider postremus a separate fronted into Cervalces with moose a single genus species of the broad-fronted moose: Cervalces two Cervalces sensu strictu and subgenera: postremus. This moose was the last form of

Libralces, and considered the true moose Alces as broad-fronted moose in Eurasia, and is either the other of Alcinae. Others consider the genus considered as the ancestor of the true moose differences in skull structure and antler features Alces (Heintz & Poplin, 1981), or as the initial between broad-fronted moose and true moose of the Cervalces form American stag-moose of insignificant and proof gradual evolutionary (Azzaroli, 1981). Guthrie (1990b: 112) consi- therefore all the changes; they join moose into dered that "sometime during the last glaciation, undivided Alces & 1981; genus (Heintz Poplin, Eurasian Cervalces latifrons underwentrapid with Kahlke, 1990; Lister, 1993). We disagree evolutionary changes toward anatomically this opinion, and considerCervalces and Alces to modern moose (Alces alces)”. However, the be different on the level. that broad-fronted the generic hypothesis moose were

ancestors of modern moose was already Giant broad-fronted moose Cervalces latifrons in an early based Northern Eurasia: rejected stage (Soergel, 1912), were widely distributed over the of evolution. upon irreversibility He argued west-ward to the British Isles and east-ward to that it is impossible to fit the shortening of the lower stream of the Kolyma river (fig. 5) antler beams from the broad-fronted moose to (Vangengeim, 1961,1977; Vereshchagin, 1967; the true moose into a single evolutionary Sher, 1971,1986; Vislobokova, 1990). In Siberia lineage, as it would mean a reversal of the the oldest remains of this moose belong to the hypertrophy developed in the former ones. end of the Pliocene or the beginning of the Furthermore, the shortening of the beam in Pleistocene. An antler fragment similar to that Alces the of is accompanied by appearance a of C. latifrons was found in Western Siberia new character: the expansion of an antler (Irtysh river, Podpusk village) in sediments palmation. Vereshchagin (1967:10), in our correlated with Upper Villafranchian (Vislo-

31 A review of the of Pliocene and Pleistocene 1 systematics moose, part

Fig 5 Finds of fossil remainsof moose in Eastern Siberia and the Russian Far East: a - broad-fronted moose

- Cervalces b - Siberianbroad-fronted C. true Alces d - northernlimits of the latifrons; moose postremus; c moose ; present-day distributionof A. americanus in Eurasia

Vondsten van fossiele resten van eland in Oost Siberië en verder naar het oosten: a - breedhoofd-eland, Cervalces

b - de Siberische Cervalces eland Alces d - latifrons; vorm, postremus; ware ; noordelijke grenzen van huidige ver- A. americanus Eurasia spreiding van in

opinion, fairly considered: hypothesis that 1987). Stag-moose had the characteristic the broad-fronted the direct Cervalces skull with nasal which moose was ancestor long bones, of the modern moose is doubtful. Most likely, contact the premaxillary bones, an antler beam the former about of and and antler one occupied in moose lineage average length massivity, an the same position, what in the proboscidean shovel divided into two or three parts. lineage - mammoth in relation to the Indian It is supposed that some facial skull and antler elephant... It is hard to believe in a transforma- shape of unclear form Alces brevirostris Kretzoi tion of high specialized long-beamed antlers of from the Middle Pleistocene of Hungary indi- thebroad-fronted moose during the first half of cate a connection between C. latifrons and A. the Pleistocene into the short-beamed antlers of alces (Vorbs, 1985, after Kahlke, 1999). the modern ". Azzaroli's moose Thus, assump- Remains of the found tion (Azzaroli, 1981) that C. postremus was true moose (Alces) are in ancestral the American Cervalces dated from the late Middle Pleistocene to stag-moose deposits scotti to most onwards. Alces finds from the seems us plausible. sp. Netherlands, Kola peninsula, southeast of Western Siberia, Remains of Cervalces, close to the Scott's stag- and the Enissey river basin belong to that found America's Middle may moose, are in North period (Erdbrink, 1954; Gromova, 1965; Pleistocene sediments. During the Late Pleisto- Vereshchagin, 1967; Alexeeva, 1980; Sher, 1986). cene C. scotti was widely distributed in this As the finds testify, in the beginning of the Late continentand existed here prior to the begin- ning of the Holocene (after Churcher & Pinsof,

32 CRANIUM 22, 2 - 2005

Pleistocene modern moose were already widely recognised: C. latifrons (Johnson 1874), Late Plio-

distributed in Eurasia. cene (Late Villafranchian) to early Middle

Pleistocene of Eurasia and Alaska (?), C. Although substantial research has been carried postremus (Vangengeim et Flerov 1965), late out on the fossil moose of Eastern Siberia and Middle Pleistocene of Eurasia, and C. scotti the Russian Far East (Vangengeim, 1961; 1977; (Lydekker 1898), Middle to end of Late Pleisto- Vangengeim & Flerov, 1965; Russanov, 1968; North America. cene, of their Sher, 1971,1986), many aspects syste- Differential diagnosis. Differs from Libralces in matics, evolutionary relations and distribution longer facial part of the skull, longer and higher solve these are still unclear. In order to ques- canines nasal aperture, absence of upper and tions, we studied material of fossil and subfossil the Paleomeryx-fold, and in theprominence of moose in various museums. We examined not nasal bones. Differs from Alces in longer and undescribed collection but only new, material, relatively thinnerantler beam (its length always also specimens described by earlier authors, exceeds the circumference), in the position and because measuring of certain antler dimensions form of the of the upperprocesses premaxillary (for example, beam length) is rather subjective in shorter bones, in a broader occipital area, and of such consequently comparison parame- rostral of the nasal and part skull, longer bones, ters becomes difficult. For this purpose we in a considerably smaller prominence between investigated material earlier described by Fisher the antlers. & Rouillier (1846) on exhibit in the Paleontolo-

gical Museum of RAS (Moscow): the so-called Cervalces latifrons Johnson, 1874 - broad-

Alces savinus G.Fisher, 1834 (Savin's moose), fronted moose and also material investigated by Pavlow (1906) Alces latifrons Johnson, 1874 Alces H. stored the as fossilis v. Meyer, 1832, in ? Cervalces alaskensis Frick, 1937

Geological Museum of RAS and some other Libralces reynoldsi Azzaroli, 1953

collection specimens. Finally, we found some fossil Alces the field more remains during expe- Material and locality. A fragment of a neurocra- ditions of the Mammoth Museum. niumwith a right antler beam, MB without

number, Adycha river, Ulakhan Sullar outcrop, found in 1978. Fragments of antlers: two speci-

FAMILY CERVIDAE GRAY, 1821 mens without numbers, MB and VMN, YIGS

4437, all from Ulakhan Sullar; YIGS 62, Aldan

Khara-Aldan river, outcrop (fig. 6a); MB with- Subfamily Alcinae Jerdon, 1874 out number, Adycha river, Oskhordokh out- Genus Cervalces Scott, 1885 found in 1994 A of the crop, (fig. 7e). fragment frontal of skull with left antler part a beam, Libralces Azzaroli, 1981 YIGS 4412, Oskhordokh (fig. 6f). Mandibles: Alces Heintz, Poplin, 1981 YIGS 4403-4405 (fig. 7a-b) and one without all from Oskhor- Type species. Cervus americanus Harlan, 1825 [= number, private collection, without UlakhanSullar. Cervalces scotti (Lydekker, 1898)], Pleistocene of dokh; number, Frag-

North America. ments of metatarsal bones: MM 8034 and VMN

without number, both from Ulakhan Sullar; MB Diagnosis. Rostral part of skull extends forward

much less than in Alces. Thewide nasal without number, Oskhordokh.

of the bones make processes premaxillary Geological age. YIGS 4403-4405, 4412 and 4437 contact with the nasal bones. Nasal bones are are found in deposits of the Adycha site, dated than the dental elongated, longer upper row. Late Pliocene - Early Pleistocene (Anonymous, The cerebral section of the skull is wideand 1987). YIGS 4403-4405 are found together with low. The ratio of cranium height from the lower remains of Early Pleistocene representatives: edge of foramen to maximal occipital magnum Archidiskodon cf. Canis sp., Trogontherium cuvieri, width is usually less than 70%. A small promi- cf. variabilis (Kotov et al., 1976). Earlier remains nence is present between the antlers. The orbits of broad-fronted moose were also found in are much lower than the roof of the forehead. these outcrops together with Early Pleistocene The antler beam of is long or average length EquusMammuthus trogontherii, (Plesippus) cf. (length always exceeds beam circumference), verae. YIGS 4437 is found together with Bison and is slightly bent. priscus andRangifer cf. tarandus. Antler frag- Species composition. At least three species are

33 A review of the systematics of Pliocene and Pleistocene moose, part 1

of 1 - skull Table 1 Sizes the neurocranium(in mm) of broad-fronted moose and stag-moose (genus Cervalces).

the lower of foramen 2 - skull from the of foramen 3 - height from edge magnum; height upper edge magnum;

maximal occipital width; 4 - width of occipital condiles; 5 - width of the skull at the constrictionbehind the ant-

- lers. Q1 - Early Pleistocene; Q2 - Middle Pleistocene; Q3 Late Pleistocene; Q4 - Holocene.

breedhoofd-eland hert-eland 1 - Matenvan de hersenschedel (in mm) van ( latifrons ) en van (geslacht Cervalces).

schedelhoogte vanaf de onderrand van het achterhoofdsgat; 2 - schedelhoogte vanaf de bovenrandvan het ach-

- maximale - breedte de - terhoofdsgat; 3 achterhoofdsbreedte; 4 van achterhoofdsknobbels; 5 breedte van de

- de achter het - - Holo- schedel ter hoogte van insnoering gewei. Q1 Vroeg Pleistoceen; Q2 LaatPleistoceen; Q4

ceen.

Cranium Cranium Cranium Cranium Cranium Cranium . , , , Index:n 6X 1:3 height 1 height 2 height 3 height 4 height 5 heightheight 6 '

Moldavia, Kolkotova Balka, Q1 (David et al.,

- 100 240 - - - - 1990) Cervalces

latifronslatifrons Ulakhan Sullar, without No, VMN 146 106 209 118 - >120 69,9

Oskhor-dokh, YIGS 4412 - - - - - 136 - j- j- j-

C. postre-postre- Yakutia, Q2 Verkhnevilyuiskoye, YIGS 1277 - - - - 125 - mus

Alaska, Q3, AMNH 342-2426 122 82,4 184 105,2 H9119 107 66,3

Alaska, Q3, AMNH 627-6546 119 81 167 103 114 >100 71,3

Alaska, Q3, AMNH 101-5090 115 72 >170 - 114 >105 <67,7

C. scotti Alaska, Q3, AMNH 712-1342 - - >170 87,3 122 - -

Alaska, Q3, AMNH 198-5899 115,4 83,1 166 100,1 112 >110 69,5

Alaska, Q3, AMNH 16250 118 70,1 182 97 113 >110 64,8

New Jersey, Q3-Q4, AMNH 107678 124 83 178 96,4 113,5 >120 69,7

of - skull from Table 2 Average sizes neurocranium (in mm) in true moose (genus Alces). Measurements: 1 height

the lower of foramen 2 - skull from the of foramen 3 - maximal edge magnum; height upper edge magnum; occipital width; 4 - width of occipital condiles; 5 - skull width at the constriction behind the antlers; 6 - distance

* from the burr to the midpoint of the frontal bone. Note: measurements from photo.

- vanaf Gemiddeldematen van de hersenschedel (in mm) van ware eland (geslacht Alces). Maten: 1 schedelhoogte

de onderrand het 2 - vanaf de bovenrand het 3 - van achterhoofdsgat; schedelhoogte van achterhoofdsgat; maxi- male achterhoofdsbreedte; 4 - breedte van de achterhoofdsknobbels; 5 - schedelbreedte ter hoogte van de insnoe-

*

het - afstand de het voorhoofdsbeen. ring achter gewei; 6 van stok tot middelpunt van Opmerking: maten

foto. genomen van

Alces ameri- Alces ameri-

Alces cfcfAlces AlcesamericanusAlcesamericanus canus, mo- canus, mo-

dern dern

Yakutia Lo- Measu- EasternFastprn Eu-Fu Yakutia Ko- Extreme ÏL. Western Si- Yakutia, Q4, . wer Chukotka, Central rements ™ werKoIy-Koly- rements Du- North-East rope, Q3.Q4,Q3-Q4, lyma, wwithout. thout beria, Q3-Q4 ma, Q3-Q4, MalyiAnyui,MalyiAnyui, Yakutia, GMGMRAS,RAS, , Yar, AM 1067527 of Siberia, /A1 vanny ™ (Alexeeva, number,number, without Q3, YIGS r/x *x t (Alexeeva, A.a.A.a.pfizen-pfizen- ZMMSU, Q3,MMMM ZMMSU, ; nom v.~ Q3, , A.a.A.a.gigas,gigas, n=2 4291 n=7n=7 . 1980)1980) YIGS, <7. ™ number, Q ' mayeri, n=4 8121 n=9 « n=2

11 121 135 126,5 JL29129 127,3 J123123 136 120,.8 125,.2125,2

2 86,2 80 84 82 83,5 85 100 _82,382,3 86,3

3 _168168 JL80*180* 173,8 162162 175,5 170 _181181 159,.6 169,8

4 101,7101,7 104*10V 9%599,5 104 105,5 99 103 96,696 6 98,6 _4 J104 J103 £

- _55 9393 109 101,5101,5 - JL02102 JL05105 105 102,4 106,1

6 - - _6 - 99,5 JL01101 99 J04104 92J92,7 100,1

Index 1:3 72,3 7575 72,872,8 79,679,6 72,5 72,4 75,1 74,8 73,3

34 CRANIUM 22, 2 - 2005

Fig 6 Fragments of antlers of the broad-frontedmoose

- YIGS - C. latifrons: a Aldan river, Khara-Aldan, 62; b, c

Adycha river, Oskhordokh, after Lazarev & Tomskaya,

1987; d - C. aff. latifrons, Bolshaya Chukochiya river, after Sher, 1971; e - Adycha river, Oskhordok, specimen without number, MB; f - skull fragment with part of antler beam, Adycha river, Oskhordok, YIGS 4412 (the area of the frontal elevation is marked with an arrow)

C. lati- Fragmenten van geweien van breedhoofd-eland, frons: a - Aldan rivier, Khara-Aldan, YIGS 62; b, c - Ady-

& cha rivier, Oskhordokh, naar Lazarev Tomskaya,

1987; d - C. aff. latifrons, Bolshaya Chukochiya rivier,

- stuk naar Sher, 1971; e Adycha rivier, Oskhordokh,

zonder f - nummer uit MB; schedelfragment met een ge- deelte van de geweistok, Adycha rivier, Oskhordokh,

de YIGS 4412 (het gebied van de opbolling aan voorzijde is aangegevenmet een pijl)

Fig 7 Mandiblesof Alcinae. C. latifrons, Adycha ment YIGS 62 from Khara-Aldan, described river, Oskhordokh: a - YIGS 4403, b - YIGS 4404; C. earlier Russanov the to - by (1968), belongs Early scotti : c AMNH 107678, New Jersey,

Pleistocene. Mount-Hermon, d - AMNH 198-5899, Alaska,

Engineer Creek; e - A. americanus, AMNH 276-283, Description and comparison. All material of C. Alaska latifrons is strongly mineralized, coloured

brown to dark - brown. Traces of Onderkaken Alcinae. C. transport by van latifrons, Adycha river, flow revealed the described water are not by Oskhordokh: a - YIGS 4403, b - YIGS 4404; C. scotti: remains. c - AMNH 107678, New Jersey, Mount-Hermon, d -

AMNH - A. The skull fragment from UlakhanSullar is 198-5899, Alaska, Engineer Creek; e AMNH Alaska represented by occipital, parietal and partial americanus, 276-283, antler beam. frontal parts with a preserved right The is On skull specimen very large (table 1). but it is noted that broad-fronted moose had a fragment YIGS 4412 part of the frontal bone larger occipital width than Alces (after Vislo- from antler basement to the sutura frontalis and bokova, 1990). The specimen from Ulakhan part of the left antler beam are preserved (fig. Sullar, which has a smaller occipital width than 6f). In YIGS 4412 and in the previous specimen from Moldavia, however, C. latifrons surpasses the from the burr to the distance (coronet) up this parameter in true moose. Broad-fronted middle of forehead (sutura frontalis) is larger differ from Alces moose reliably in occiput (table 1) than in true moose Alces (table 2). For height and in width of the occipital condyles broad-fronted moose the small elevation at the (tables 1 and 2), in mandible size (tables 3 and 4) middle of the forehead is characteristic (fig. 6f). and in teeth (tables 5 and 6). It is obvious that C. Skull remains of C. latifrons are poorly known,

35 A review of the systematics of Pliocene and Pleistocene moose, part 1

section is and wide. latifrons had a much larger skull than Pleisto- The symphyseal high The molars have additional forma- cene and modernAlces. The lower molars of C. strengthening latifrons, too, are much larger than those of Alces tions, counteracting wear. The second type of (tables 5 and 6). Having undoubtedly larger lower jaw constitution is characteristic for forest

than the C. bisons, mild forest branches, jaws true moose, latifrons preserved eating grasses, elon- leafs and barks of and bushes. For them some original proportions: a relatively trees gated dentition (about 39%) than Alces (33-37%), the low dental section and molars without and shorter formations higher jaw and dental sections, a strengthening are peculiar; symphy- diastema (fig. 7; tables 3 and 4). The lower seal section is narrowed and low.". In our molars of the broad-fronted moose always have opinion, a similar division in lower jaw types is labial pillars, increasing the masticatory surface. seen in broad-fronted and true moose. Cervalces,

in These pillars are not always present Alces. inhabitant of open landscapes, apparently and also leafs These specific features allowed the broad- mainly ate rigid steppe grasses, fronted moose to chew more rigid nutrition. and branches of bushes. True moose obviously

It is interesting to note that Flerov (1979: 22) were always more connected with forests and and and noted in bisons two similar types of lower jaw bushes, branches water-paludous "The inhabitants in their diet; and the herba- constitutions (in translation): green prevailed

whose ceous remained minor food The of open steppe landscapes, feeding plants a source. consists of rough forage (mainly grasses), are specalization of feeding behaviour of true dental of the of the facial characterized by high section moose promoted lengthening lower jaw, and are thus capable to maintaina section of the skull during evolution. Lengthe- of foremost of the skull and the large pressure while grinding firm vegetation. ning the section

Table 3 Sizes of mandibles (in mm) in broad-frontedmoose and stag-moose (genus Cervalces). Length of 1 - man-

- - - - diastema. - from the dible; 2 tooth row, coronar; 3 same, alveolar; 4 premolar row; 5 molar row; 6 Height 7

8 - ahead P 9 - under the middle of 10 - behind M3. coronar process; 2; M1;

hert-eland Cerualce 1 - Maten van onderkaken (in mm) van breedhoofd-eland (latifrons) en (geslacht s). Lengte van

- - - 2 - de 3 - de 4 valse 5 6 onderkaak; tandenrij, over kronen; idem, langs tandkassen; rij kiezen; rij ware kiezen;

diasteem. Hoogte 7 - vanaf coronair uitsteeksel; 8 - net vóórP2; 9 - net onder midden van M1; 10 - net achter M3.

Cervalces latifrons C. cf. latifrons C. sp.,Q3 C. scotti, Q3-Q4

Yakutia, Western Sibe-Sibe- i, , . iUlakhann TIransbai-Transbai- Yakutia, Adycha river, Oskhordokh, . . i AlaskaA1 i Alaska XT T ~ ria, Ob river, Alaska Alaska New „ , -. ria, river, JerseyJersey Q1-Q2 Sullar, kalia,kalia, Q1 ni n9 WQ1-Q2 Q1-Q2

without without Kozham-

YIGS YIGS YIGS numb er ™" number,' (A1(Alexee- Unf Unf Unf number, kulova, AM 1085745 AM 1985899 AM 107678 t 44034403 4404 4405 private col- va,1980) VMN 1974 lection

- 1 - 510 - - 485 - 460460 400 442 j

2 196 198 - 186 - - 177,5177,5 179 JL52152 162

- 3 201201 202 - - 204204 190 183 154154 164 j- J183

4 8888 88 85 82 - 84* 76,576,5 75 6363 73,5

5 111111 108 - 104 - 109* 103,2103,2 105,.5 89 95 _5 J104 j-

- 6 -- 185 - - - - 174 144 163 j JL63

7 - 251 - - 257 - - 226 223 221 j

- 8 49,3 51 J5555 55,5 - - 43 43 48

9 64 65 - - 64.2 - - 53 59 60,3 J>5 j _59

10 80 85,5 - - - - 72,972,9 71,4 7373 74,7 j j j

Indices:

71 4:5 79,3 81,5 - 78,9 - 77,1 72,072,0 71 _70870,8 77,577Jb_

2d2:1 - 38,8 - - near 40 - - 38,9 38 36,7 j

6116:1 - 36,3 - - - - - 37#37,8 36 36,9 j

7:1 - 49,2 - - near 50 - - 49,1 near 50 50

36 CRANIUM 22, 2 - 2005

diastema promoted increase of mouth cavity broad-fronted moose and of Alces the fifth and, accordingly, has enabled them to improve molarization degree after Heintz is characte- of the for fused with and manipulation tongue grabbing ristic: protoconid is entoconid, branches and twigs (Gogina, 1979; after Vislo- hypoconid is separated (after Vislobokova, bokova, 1990). 1990).

In various lines of deer an evolutionary shorte- Antler beams of C. latifrons from Yakutia and

of the is observed. At the from other of the ning premolar row parts geographic range are

same time the premolar row remains rather similar in size and proportions (table 7). Broad-

in found fronted has antlers 2 long moose (Vislobokova, 1990). We a moose very large (spread rather high index of premolar length in C. lati- to 2,5 m) with a long beam (apparently, up to and modern Alces. At the The index of their frons same time, it 0,6 m) (fig. 4a). massivity

to be lower in fossil and antlers is 42 to the circumference of appears slightly 82%, i.e., an subfossil Alces (table 4). Premolars are mola- antler beam is always less than its length rized C. well Alces: the P3 where this index is than in latifrons, as as in (contrary to Alces, more valley between paraconid and metaconid is 100%). Similar antler dimensions and propor- closed. of broad-fronted characteristic for Pleistocene A P3 an archaic moose tions are Early C. is described from the river Bolshaya latifrons from the huge territory of Northern

which has from Western to Chukochiya, a deep valley between Eurasia, ranging Europe up these conids (Nikolsky, 1996). For the P4 of Eastern Siberia (table 7). The antler beam of this

4 Sizes in 1 - mandi- Table of mandibles (in mm) and proportion indexes (%) true moose (genus Alces). Length of

ble; 2 - tooth row, coronair; 3 - tooth row, alveolar; 4 - premolar row; 5 - molarrow; 6 - diastema.Height of the

mandible 7 - from the 8 - ahead P2; 9 - under the middleof 10 - be- coronair processes; height height M1; height

hind M3.

Maten (in mm) en verhoudingen (in %) van onderkaken van ware eland (geslacht Alces). Lengte van 1 - onder- kaak; 2 - tandenrij, over de kronen; 3 - tandenrij, langs de tandkassen; 4 - rij valse kiezen; 5 - rij ware kiezen; 6 -

diasteem. - vanaf het 8 9 - onder midden 10 - achter Hoogte 7 coronair uitsteeksel; net vóór P2; net van M1; net M3.

Alces.A/ces. cf.cf. A. americanus, Yakutia Q3 AlcesAlces alces, americanus,americanus, A. americanusAlaska, Q3-Q4 A. americanus,americanus,, modern -Q4 modern Yakutia, Q3

Olyek- Ady- Koly- ma ccha,ha, v , Koly-woty- ma, basin, Osk-Dek. LenaT ma AM AM

AM ... AM AM AM ... . Eastern North-East , ... 04 1000- ma, Q4,v c ' Western hor-hor- coo AM Western Si- delta, _ . 583 4854 27652 5832 36814 of Q3- wit- 15000 5842 Siberia, Siberia, dokhdokh YIGS 616 beria, n=9 Q4/MMQ4,MM hout 734 83 162 26 n-14 n-12 hout yearsyears 2167 161 4759 7099 num- B.P., YIGS ber MAE, 44II4411 n=3n=3

-- 1 - - 472 492 485 470 482482 520 502 501 510510 453,8453,8 468,4468,4 486,4

2 180 167 155,5135,5 163,2 160,7 171 - 173173 157 169 168 177177 159,1159,1 161,1 167,5

33 _7474 171 159 168 162,6 174 167 175175 161 172 172 179 - 164,7164,7 168

4 100 70,5 69 _7373 69,2.69,2 69,5 70,5 74,574,5 68 _68_68 __6969 74 66A66,4 68,568,5 69,6

55 - 100100 91 94 94,294,2 9898 96 101101 93 84 99 103 81 81,881,8 88,3

66 - 170 199 201 - 205205 190190 193193 224 210 208 202 - 181181 192 j-

77 49 - 210 226 -- 219 215 234234 238 236 237 240 211,4 213,2213,2 220,7

_88 59,5 44 47 53 -- 55 4949 54 54,3 58 53 52 - 47,1 49,749,7

60,2

9 - - 64,5 63 - 58 50,5 63 66,8 63 62 60 - 56,256,2

JtO10 - 67 72 - 72 60,5 73,2 79 72_72 70 71 - 66 70,3

4:5 74 70,4 75,7 77,7 73,2 70,8 73,4 73,8 73,1 81 jil81 71,9 82,7 82,9 80,380,3

2:1 - - 32,9 33,2 - 35,3 35,5 35,9 30,2 33,7 33,533,5 34734,7 _35335,3 35,1 35,235,2

6:1 - - 42,2 40,9 - 42,3 40,4 40 43,143,1 41,8 41,5 39,6 - 38,7 39,139, 1 j £2,2

- 7:1 - - 44,544,5 45,945,9 - 45,2 45,8 48,5 45,845,8 47 47,3 47,1 45,9 45,3 45,145,1

37 of the of Pliocene and Pleistocene 1 A review systematics moose, part

Libralces whichbroad-fronted be moose is slightly curved, in contrast to moose can disting- gallicus, which has an S-shaped beam. Sher uished from Alces. (1971) describes an antler fragment of broad- broad-fronted Remarks on systematics. The fronted moose from the Olyerian deposits at the moose remains from Yakutia investigated by us river Bolshaya Chukochiya, which has a more does practically not differ from the earlier thin and curved beam than C. latifrons (fig. 6d; described C. latifrons remains from Europe and table 7). This observation, and also the weak Siberia. molarization degree of the P3 of the Olyerian The original feature of Libralces, Cervalces lati- allows archaic moose, to consider it as an form, noted frons and C. scotti, as it was already probably intermediate between L. gallicus and the nasal which above, are long bones, come C. latifrons (Sher, 1971; 1986; Nikolsky, 1984). of the into contact with the upperprocesses The antler fragment fromOskhordokh (fig. 6e; premaxillary bones (Azzaroli, 1952,1981; Sher, table 7), too, has a bent beam. 1986) (fig. 3a-c; fig. 9d-f). This feature puts them Metatarsals of broad-fronted moose are large together with deer of the subfamily Cervinae: and slender. Metatarsals of C. latifrons are much Cervus, Megaceros, Rangifer, Capreolus, etc. (fig. larger than those of Alces, but have similar 3d; fig. 9c), and also distinguishes them from proportions (fig. 8, tables 8 and 9). Sher (1971) the the true moose Alces (fig. 3e-h; fig. 9d-f). At noted some metatarsal features on the basis of same time broad-fronted moose and true moose

of lower teeth Table 5 Sizes (in mm) jaw of the broad-fronted moose and stag-moose (genus Cervalces)

hert-eland Maten (in mm) van onderkaakskiezen van de eland met breed voorhoofd(latifrons) en van (geslacht Cervalces)

P M P2P2 P33 P4 MiMI M22 M3MJ

length width length width length width length width length width length width

Cervalces latifrons

Western Siberia, Villaf-

ranchian (?)(?) (Alexeeva 2727 17,3 29,8 22 - - - - ...- - - - 1980)

Transbaikalia,Q1

------34 25 48 >26 (Kozhamkulova 1974)

Yakutia, Adycha, Osk- 24 18,5 31,1 21 32 23 33 2424 37 25 48 23,3 hordokh,Ql(?),YIGS 44034403

46 YIGSYIGS44044404 23.223,2 - 29,4 21 33,3 25,2 31 26,126,1 35,7 27,2 46 27,1

- - - YIGS4405 - - - 21.321,3 14,8 30,2 19,7 36,5 23,7 _-

Yakutia, Adycha, Ulak-

han Sullar, Ql(?), MB ------47 23,5

n.n..

C. cf. latifrons

Western Siberia, Ql- 20,1 13,1 23,6 17,5 28,5 20,3 28,05 22,822,8 30 24,8 45,5 24,3 Q2(?) (Alexeeva 1980),1980),

Q2-Q3(?), Alaska, 2222 15 29 20,1 35 24 34 24 37 25,4 46 23,4 AMNH 527-4102

C. aff. latifrons

Yakutia, Bolshaya

Chukochiya, Late Villaf- - - 29 19 34 2323 - - 36 24 - - rancian (Nikolsky 1996)

C. postremus(l)postremus (?)

Adycha, Oskhordokh,

------43,6 22,5 Q2(?): YIG54406YIGS4406

------44 YIGS44364436 - 36,3 23,2 44 24,5

C. scotti

New Jersey, AMNH 21,721,7 15,5 28 18,4 30,6 20,520,5 30,2 20,720,7 31 21,1 41,3 19,5 107678

Alaska, AMNH 1985899 1919 12,3 23,4 16,6 26,8 18,8 28,2 21,4 30,2 20,5 39 20,1

38 CRANIUM 22, 2 - 2005

which them unites the with not share many general features, unite Caucasian moose Cervalces,

into a uniform subfamily Alcinae and disting- with Alces. For Lister this observation proves

uish them from Cervinae: a horizontal location the affiliation of the broad-fronted and true of antler antler wide forehead Alces. At close considera- beams, shape, moose into one genus

that is inflated in the frontal sutura region, tion it also becomes clear that the short rostral

choanalheight, constitution of the temporal section, as well as the nasal and premaxillary region, telemetacarpality, etc. (after Vislo- bones of Cervalces are extremely similar to those

bokova, 1990). The name Cervalces ('deer- of the Cervinae (fig. 3d; fig. 9c). For all Alces a

-moose') is appropiate, since it reflects features lengthened rostral section and short nasal bones

In of the of both moose and deer. are characteristic. addition, the shape

nasal bones in Alces differs completely from that One of the basic features of Cervalces (the combi- Cervalces and deer. The of in upper processes nation of long nasal bones, which come into the premaxillary bones in Alces are long and contact with the of the upper processes thin, with an expansion at the end in the Euro- premaxillary bones) is never found in Alces. The features pean moose (fig. lOa-d). specified Sher (1986) and Lister (1993) demonstrated that of skull of the rostral part the modern moose subfossil Caucasian moose A. alces caucasicus are unique. has and wide of the long very upper processes premaxillary bones, which come into contact "All these features lead to a significant increase with the nasal bones (fig. 3g; fig. 9e); this feature of the dimensions of an osteal nasal foramen

Table 6 Sizes (in mm) of lower jaw teeth of the fossil and subfossil true moose (genus Alces)

eland Maten (in mm) van onderkaakskiezen van fossiele en subfossiele ware (geslacht Alces)

MlMi M2M P:P2 P3 P4 2 M3Ms

length width length width length width lengthlength width length width length width

Alces cf.ei. alces,alees, Western Siberia, Q3- 17,5 13 22,9 13,3 27,5 20 25,8 21,5 28,1 22,922,9 40,1 22,8 Q4 (Alexeeva, 1980), mean.

A. A. sp., Yakutia, Q3(?)

Adycha, Oskhordokh, YIGS 4411 21 16 27,1 20 28,7 2121 28 21,2 31,3 21,4 - -

Lena delta, YIGS 4759. 17,5 13,6 25,2 17 [2929 19,419,4 28,3 20,2 31,5 22 43,3 22

A. cf.et. americanus,americanus, Q3

Adycha, Ulakhan Sullar, YIGS 4435 - - 27,4 17 - - - - 30,5 21,7 40,2 21,521,5

Sullar, YIGS 4959 - - 25 16,8 28 19,5 28 20 29 20,4 40 19,4 Adycha, Kyra j-

Rebrovo site, no number ------31,4 24,3 - - - - j- j j- j- j-

YIGS 5614 ------38,5 16,4 Alazeya, j-

no number,MEYUMGO - - 23 18 28,7 21 ------Ynykchan, j- j- _18

Kolyma, MM 7099. 18,7 14,5 23,4 18,6 2828 19,8 28,1 20,9 32,8 23,6 40,3 22,8

A. americanus, Q4

Olyekma river, Ulakhan Segelen------32,3 24,2 42,7 21,5 nyakh, Neolithic

Olyekma river, Ulakhan Segelen- nyakh, Early Iron Age, MAE, 18,1 13,8 23,7 17 29,1 19,5 26,1 20,5 29,5 20,9 40,6 20,8 1000-1500 y.B.P.,y.B.R, MAE

Kolyma, no number 19.519,5 13,2 24,3 17,5 28 21 26,3 20,2 30 24 41,5 22,2

A. americanus, Alaska, Q3-Q4

AMNH 583-2167 JL818 13 25 18,5 29 21,2 29 22,5 32,2 23 44,7 21,6

AMNH 616-6015 21.621,6 16 25,4 20 30 23,2 27,2 22 _3131 _2323 47347,1 22

AMNH 276-5283 1818 13 24,2 18 28,5 20,7 28 22 31,5 23,4 43,2 22,3

AMNH 276-5285 19,6 15,6 25 _2020 30,2 20,8 29,6 22,322,3 33,3 22,1 44,2 23

AMNH 473-3436 20,1 15,6 24,3 18 28,4 20,4 31,1 2323 32 24,2 45,4 23

AMNH 584-2161 19,9 15,5 2323 17 28,5 20 28 22 30,7 23 43,8 21

39 A review of the systematics of Plioceneand Pleistocene moose, part 1

a b c

9 of fromYakutia: - C. b - Alces Fig Metatarsal bones moose a latifrons, Adycha river, Oskhordokh; sp., Adycha

- Alces Lazarev & river, Ulakhan-Sullar; c sp., Sartang river (after Tomskaya, 1987)

- - b Alces Middenvoetsbeenderen van eland uit Yakutië: 1 C. latifrons, Adycha rivier, Oskhordokh; sp., Adycha

- Alces rivier Lazarev & rivier, Ulakhan-Sullar; c sp., Sartang (naar Tomskaya, 1987)

Table 7 Sizes (in mm) and proportion (in %) of antler beam of the broad-frontedmoose. 1 - beam length; 2 - cir-

- cumference of burr; 3 - circumference of beam above the burr; 4 horizontal diameterof beam

de eland breed voorhoofd 1 - Maten (in mm) en verhouding (in %) van geweistangen van met (latifrons ). lengte

- - de - omtrek de 3 omtrek de 4 van stang; 2 van rozenkrans; van stang boven de rozenkrans; stangdoorsnede

Cervalces

aff. lati- Cervalces latifrons, Europeanpart and Ural Cervalces latifrons, Eastern Siberia frons

Aldan

Ger- river, Bolshaya Ger- FEastern8"1 Oskhor- Ulak- Kha- Chukoch Ti- , Adycha Adycha river, many, Moldavia, c river, Ural Siberia,w, many, ' Sizes T/T Sussen-Süssen- dokh, Q1Ql hanSullar,Q1Ql ra-Al- iyaiya river, u raspol (?) MMosbach° sbach Q1Ql bornborn dan, Q1Ql

QUQl.

Lazarev " Kahlke,ablke YereshcVereshc YakhiYakhi- ' Vangen- f Kahlke, David, & yiGSYIGS MB Y1GSYIGS VMN MB Y1GSYIGS Sher, 1956, hagin, movich, geim, 19561956 1965 4412 n.n. 44374437 n.n. n.n. 62 1971 mfv"' ?Q1T' Tomskay1 1965 1961 1960 1967 _ 1QQ a, IVo/1987

11 382 354354 340 440 429 350 - 280 550 250 390 365 440 520

- - 22 300 282282 351 - 222 313,3 307,5 312 330 345 340 305

33 223 223223 280 224 180 243,3 121,5 230 275 - 250 230 254 220

44 67 70 94 - 62 - 68 70 - _5757 - - 74 J>060

3:13:1 57,9 69,269,2 82,3 51 42 78 73,4 82 50 - 64 63 57,7 42,3

40 CRANIUM 22, 2 - 2005

Fig 10 Skulls in top view: a - C. latifrons, Süssenborn; b - C. scotti, New Jersey, Mount-Hermon(both after Sher,

1986); c - Cervus elaphus (after Sokolov, 1979); d - Alces alces; e - A. alces caucasicus (after Vereshchagin, 1949); f - A. americanus

onderschrift Schedels gezien van boven; voor soortnamen en vindplaats, zie Engelstalig

and indices and Table8 Measurements(in mm) (in %) of metatarsal bones of broad-fronted moose stag-moose

(genus Cervalces). 1 - length; 2 - width of proximal end; 3 - transversal diameterof proximal end; 4 - width of dis- tal end above trochlea; 5 - transversal diameterof distal end; 6 - width of shaft at midpoint.

Maten middenvoetsbeenderen de eland breed voorhoofd (in mm) en verhoudingen (in %) van van met (latifrons)

Cervalce - - breedte 3 - dwarsdoorsnede door bo- en hert-eland(geslacht s). 1 lengte; 2 aan bovenuiteinde; venuiteinde; 4 - breedte van onderuiteinde; 5 - dwarsdoorsnede door onderuiteindeboven de rollers; 6 - breedte van de schacht in het midden

CervalceCervalcess C. cf.postpostre-re- C. scotti, Cervalces latifrons, Q1Ql aff. latifrons, C. cf. latifrons, Ql mus,mits, Q2 Q3 Q1Ql

Measu- Adycha Bolshaya New JJer-er rements Yana ?,?' Alaska TiraspolF Ulakhan Sul-Sul S^ MalyAnyuiMaly' Anyui' Oskhordokh Chukochiya sey larlar

Lazarev MM & AM AM AM Lazarev & MB VMN Lazarev , bner,Sher, iy/I1Q711971 Sher,eh 1Q711971 Sher, 11971cj7 ' 3731709 ' nn.n.n nn.n.n 8034 1987 3731709 47446494744649 107678 Tomskaya,1987Tomskayal987 , . . . Tomskaya,

1 464 475,5 445 - - ca. 500 ca. 500 468 - ca. 460 448

- - - 64 71 67 - 2 ca. 64 59,7 64 72,572,5 62,4

3 74 67,7 - - - 72 74,5 72 - 72 64,7

4 92.742,7 76,9 84 93,6 93 - 89J89,7 76 95,3 65 _76_76

46 41II 51,6 51 50 46,3 39,5 J1646 j4949 44 37,1

47 6 56.856,8 5050 - - 54 - ca. 6060 59,3 ca. 45 48,1 j-

Indices

- 2:1 ca. 14 12,512,5 - . - ca.ca.1313 ca. 14,214,2 14,3 15,7 13,913,9 j-

- - 4:1 20 16,116,1 16,116,1 - - - ca. 17,9 16,2 17 j

6:1 10 8,6 8,6 - - ca. 9 ca. 8 9,8 - 9,6 8,3

41 A review of the of Pliocene and Pleistocene 1 systematics moose, part

which has no analogue in others cervids, and Cervalces cf. latifrons they are connected with the existence of a Cervalces alaskensis Frick, 1937 powerful system of nasal cartilages and a long Material and Collections of AMNH, in the moose" locality. mobile (proboscis-like) upper lip Alaska. Antler beams: AMNH 30496 (holotype (after Sher, 1986:13). The rostral section of Alces of Cervalces alaskensis), AMNH 30497, AMNH is somewhat similar to that of the saiga-ante- 365-4327and AMNH 710-1491, vicinity of Fair- lope Saiga tatarica, for which the ventricose banks; AMNH 527-4127 AMNH is (fig. llb-d), proboscis-like muzzle an adaptation to a 527-4128 and AMNH 433-3858, Cripple Creek. habitat in boreal regions with a continental Metatarsals: AMNH 373-1709, Ester Creek; climate (Sokolov & Zhirnov, 1998). Probably a AMNH 474-4649, Livenhood Creek. similar constitution of the front of the moose muzzle indicates its origin in the boreal part of Geological age. The specified antler beams are Eurasia. The shape of the of the upper processes referred to in the AMNH catalogue to Rancho-

bones of modern Alces was premaxillary labrean, that corresponds to the American discussed above. The of the Cauca- morphotype geochronological scale with 480 -10 thousand sian in which the nasal bones moose come into BP Churcher & years (Repenning, 1980; Pinsof, contact with the bones does not premaxillary or 300 -10 thousand BP 1987) years (after unite this moose with Cervalces at all, because Martin & Steadman, 1999), so it includes part of there essential of the nasal is no lengthening the Middle Pleistocene and the Late Pleistocene. bones in this case, and much the of more, shape All materials are strongly mineralized and the nasal bones is for Alces. Also the typical in colour from almost range black up to brown. of the bones of upper processes premaxillary Description and comparison. The beams are the Caucasian moose differ from those of the and massive their dimen- very long (table 10), broad-fronted moose, stag-moose and deer. A sions are close to those of Eurasian C. latifrons similar morphotype as that of A. a. caucasicus is (table are bent met modernA. alces: 7). They slightly (fig. llb-d), occasionally among among especially AMNH 30497, which beam is practi- 504 investigated skulls of this species we found cally S-shaped (fig. 11c). two similar specimens (fig. 10a). The dimensions of metatarsals from Alaska

Table - - 9 Sizes (in mm) and indices (in %) of metatarsal bones of true moose (genus Alces). 1 length; 2 width of

- - proximal end; 3 transversal diameterof proximal end; 4 widthof distal end above trochlea; 5 - transversal dia-

meter of distal end; 6 - width of shaft at midpoint.

Maten eland 1 - 2 - (in mm) en verhoudingen (in %) van middenvoetsbeenderenvan ware (geslacht Alces). lengte;

breedte - - van bovenuiteinde; 3 dwarsdoorsnede door bovenuiteinde; 4 breedte van onderuiteindeboven de rol-

- lers; 5 dwarsdoorsnede door onderuiteinde; 6 - breedte van de schacht in het midden.

A.a.pfizen-A.a. pfizen- AlcesA/ces A. americanus sp. A. americanus A.a.gigasA.a. gigas mayeri

Yakutia, LowerLower Tun- Yakutia, Q3 , Extreme X/ Measure-„ . . . . Central . „ KT river, Alaska, North-E„,North-East guska UlakhariUlakhan Sul- Alaska, QSk*Q3-Q4 ments Yakutia river Q3gska„v„. Sartang Siberia lar

AM AM AM AM Sto.»71Sher, 1971 n=6 n=7 Lazarev & Tomsk.ya,Tomskaya, 1987 XjlO modern, "~6 modern,°"*ll8 'T'' "'7 34586 3681426 3681426/1£$UX/1 5862310

J1 410 422,6422,6 418,3 433 420 422 429 391 410,3

2 49 _5577 54354,5 52 61 59 58 50,6 56,4

_33 54 55255,7 57257,2 _5757 63 J5959 63 55,77 59,3 _55£

4 65 70 64,8 68 73 71 73 64364,3 69

_55 39239,5 44 4343 44 44 46 _5050 42,2 45,5

_66 34334,2 33,7 31,4 34 36 36 37 32,4 33,7

Indices

2:1 11,9 J13,413,4 13,0 12 14,5 14 13,5 13,7 14,1

4014:1 J15315,9 JL6316,3 JL515,4,4 15,7 17,4 16,8 17 16216,5 17,1

6:1 8,4 [727,9 [727,5 7,9 8,6 8,58,5 8,6 8,2 8,3

42 CRANIUM 22, 2 - 2005

of the of the bone Alces - Fig 11 Shape upper process premaxillary in : a ZM 66317, Pechoro-Ilych Nature Reserve,

coll. ZM MSU, Russia; b - MS 35551, Eastern Germany (?), coll. MS, Germany; c - ZIN 33726, St Petersburg dis-

trict, coll. ZIN RAS, Russia; d - ZM 35273, Western Siberia, Malaya Sosva river, coll. ZM MSU, Russia; e - MS 494,

- Western coll. f coll. - ZIN Ukraine, MS, Germany; no number, Norway, KM, Germany; g 32700, Canada,

Ontario, coll. ZIN RAS, Russia; h - ZIN 12834, Eastern Siberia, Aldan river, coll. ZIN RAS, Russia

Vorm van het bovenste uiteinde van het premaxillare in Alces: a - ZM 66317, Pechoro-Ilych Natuurreservaat, coll.

b - MS Oost Duitsland coll. - ZM MSU, Rusland; 35551, (?), MS, Duitsland; c ZIN 33726, St Petersburg district,

- coll. ZIN d ZM West Sosva coll. ZM - RAS, Rusland; 35273, Siberië, Malaya rivier, MSU, Rusland; e MS 494,

coll. f - West Ukraïene, MS, Duitsland; - zonder nummer, Noorwegen, coll. KM, Duitsland; g ZIN 32700, Canada,

Ontario, coll. ZIN RAS, Rusland; h - ZIN 12834, Oost Siberië, Aldan rivier, coll. ZIN RAS, Rusland

Table 10 Sizes (in mm) and massivity index (=3:1, in %) of antler beams of Cervalces cf. latifrons from Alaska. 1 -

length; 2 - circumference of burr; 3 - circumference above the burr; 4 - circumference at midpoint; 5 - horizontal

diameter.

Maten massiviteitsindex in - (in mm) en (=3:1, %) van geweistangen van Cervalces cf. latifrons uit Alaska. 1 lengte;

2 - omtrek rozenkrans; 3 - omtrek boven rozenkrans; 4 - omtrek halverwege; 5 - dwarsdoorsnede.

Measurements AM 30496, holotype of "Cerualces“Cervalces AM 30497 AM 3654327 AM 7101491 AM 5274127 AM 5274128 AM 4333858 andand index alaskensis”alaskensis" Frick, 1937

J.1 380 405 320 JS70370 340 260 320

2 320 250 275 230 J300300 280 340

3 241 228 241 224 _249249 252 244

4 209 193 224 197 224 _222222 238

_55 68 56 75 65 69 72 76,3

313:1 63,4 56,3 - 60$60,5 73,2 - 76,3

43 of the of Pliocene and Pleistocene 1 A review systematics moose, part

exceed those of the true moose and of C. scotti, of Scott's stag-moose and another, larger species of and come close to those of the broad-fronted Cervalces.

from moose Eurasia (table 8). Cervalces postremus Vangengeim et Flerov,

Remarks. Remains of a moose close to C. lati- 1965 - Siberian (?) broad-fronted moose frons were found in the Yukon river basin in Material and locality. A skull fragment with Early Pleistocene fauna of Old Crow, which is part of an antler beam, YIGS1277, Vilyui river, similar to the Olyerian fauna (Harington, 1978 Verkhnevilyuiskoye locality, YIGS (fig. 12). in Sher, 1992). Frick (1937), when describing a Antler fragments: YIGS 45 (fig. 13c) and YIGS from Alaska new species of stag-moose (”C. 64, Aldan river, Mamontova Gora locality; YIGS alaskensis”), noted that the beam dimensions of 135, Aldan river, Rossypnoye locality (fig. 13b); this form are much larger than those of Scott's YIGS 970,1310 (fig. 13d-e) and 1382, Vilyui of Cervalces stag-moose. Antler remains a large river, Verkhnevilyuiskoye locality; YIGS 1762, from Alaska were under consideration and later Bolshaya Chukochiya; YIGS 4758, the delta of it was supposed that they belonged to C. lati- Lena river (fig. 13f); YIGS 6829, northern frons or a close form (Kurten & Anderson, 1980; Yakutia (fig. 14b); VMN specimen without Sher, 1971). Sher (1986) assumed that broad- number, Adycha river, Ulakhan Sullar (fig. 14a). fronted moose with long and bent antler beams Mandible fragments YIGS 4406 and 4436, from the Kolyma basin and Alaska are closer to Oskhordokh, YIGS. the subgenus Libralce s and belong to separate The studied of YIGS, species. If this turns out to be valid, the name Geological age. specimens No 45 and No 64 from Mamontova and “alaskensis” has priority by rule, and will be Gora, described re-installed. Apparently, remains of Alcinae No 135 from Rossypnoe were earlier Russanov He referred the material from North America may revail useful new by (1968).

information. the absolute of from Mamontova Gora to the late Middle For example, age and YIGS to the of one antler beam from Alaska (the Old Crow Pleistocene, 135, beginning

the Late Pleistocene Kazantsevo river), which parameters are close to those of C. (the Intergla-

+ The latifrons, was dated 33.800 2.000 y. BP (Kurten cial). locality Verkhnevilyuiskoe yielded YIGS, Nos 970,1277,1310, and 1382. & Anderson, 1980). In our opinion, this fact can specimens of theMiddle and Pleistocene testify the contemporaneous existence in Alaska Deposits Upper

Table 11 Sizes (in mm) and massivity index (=3:1, in %) of antler beams of Cervalces postremus and C. aff. postre-

The antler from Ulakhan Sullaris the Museum of the other antlers YIGS. From mus. at Verkhoyansk Nature; at

Russanov (1968) the averages are taken. 1 - length; 2 - circumferenceof burr; 3 - circumferenceabove the burr; 4 -

circumferenceat midpoint; 5 - horizontal diameter.Holotype described and measured by Vangengeim & Flerov

(1965).

Maten massiviteitsindex in Cervalces C. aff. tremus. (in mm) en (=3:1, %) van geweistangen van postremus en pos Het Ulakhan Sullarbevindt zich in het de andere in YIGS. Uit gewei van Verkhoyansk Natuurmuseum, geweien

Russanov de waardes 1 - 2 - omtrek 3 - omtrek boven de (1968) zijn gemiddelde genomen. lengte; rozenkrans; ro-

- omtrek - dwarsdoorsnede. zenkrans; 4 halverwege; 5 Het holotype is beschreven en gemeten door Vangengeim & Flerov (1965).

Yakutia

Aldan river Adycha Nort- .... . Lena Nort- . Bolshaya Kamchatka ,7,1 -1 J KamchatkaiNamcnatKa Viluv Viluy river, Verkhnevily- . . river, . ,. Ros- J Verkhnevily-J river, river hern . . T n i , Chukochiya Ulakhan J ,, , Mamontova Gora uiskoyeJ , 1t , • syp- delta Yakutia river Russa- „ Sullar noye nov,

1968 YIGS YIGS YIGS YIGS YIGS YIGS YIGS YIGS YIGS YIGS , Vaskovsky, HolotypeH YIGS 1762 45 64 135 970 1277 1310 1382 4758 n.n. n.n. 1966

1 294,6 280 225 265 190 185 173 215 205 220 185 180 270 278

- 2 213,8 235 195 290 - 230 195 - - - 206 205 260 -

3 174,2 160 155 198 150150 147 140 160 184 155 156 146 1170170 J157157

4 - - 144 161 143143 140 142 154 173 153 143 130 157 -

44 47 5 53,5 50 42 49 45 43 44 46 54 44 46 - 47 54

3:1 60,8 57,2 68,9 74,7 78,9 79,5 70,8 74,4 89,8 70,5 84,3 1 81481,1 63 56,5

44 CRANIUM 22, 2 - 2005

Fig 12 Fragments of Alcinae antlers from Alaska. A. americanus: a - AMNH 252-1939, Eldorado Creek (for compa-

- - of d rison). Cervalces cf. latifrons: b - AMNH 527-4127, Cripple Creek, c AMNH 30497, vicinity Fairbanks,

of Fairbanks. C. scotti and C. cf. scotti: - AMNH AMNH 30496, holotype of “C. alaskensis”, vicinity e 276-6360,

- - f - AMNH AMNH 235-8130, Creek, h AMNH 515-5170, Banks-at-fox, 276-8309, Engineer Creek, g Engineer vicinity of Fairbanks

Alcinae Alaska. A. americanus: a - AMNH 252-1939, Eldorado Creek Fragmenten van geweien uit (ter vergelijk).

- - - d AMNH Cervalces cf. latifrons: b AM 527-4127, Cripple Creek, c AMNH 30497, omgeving van Fairbanks,

C. scotti cf. - AMNH 276-6360, 30496, holotype van “C. alaskensis”, omgeving van Fairbanks. en C. scotti: e

- - h AMNH 515-5170, Banks-at-fox, f - AMNH 276-8309, Engineer Creek, g AMNH 235-8130, Engineer Creek,

omgeving van Fairbanks

this site The thin 23 - beam exceeds its are exposed at (Lazarev, 1980). vely (figs. 25): length

to the late The index of the beam fossil remains in question may belong circumference. massivity

amounts to 50 to 90%, which is Middle or the beginning of the Late Pleistocene in C. postremus

with of similar values for C. True moose of due to the co-occurrence remains to latifrons. Alces have shorter and Mammuthus primigenius Blumof the early type, the genus relatively

Bison priscus Bojanus, Equus orientalis Russanov, thicker antler beams (fig. 14) (Boeskorov, 1998), and E. lenensis Russanov. Antler beam YIGS the massivity index of the beam always exceeds 1762 from the Bolshaya Chukochya River, 100% (Sher, 1986; Boeskorov, 2001). Measure- found together with remains of M. primigenius ments of antler beams are listed in table 11. C.

of the and E. is has a thin antler beam of moderate early type, orientalis, most likely postremus attributable to the late Middle Pleistocene. length, contrary to C. latifrons, which has longer

the definition of other antlers is and much more robust beams Though age (table 7) (Vangen- Flerov problematic, most remains of C. postremus from geim & 1965; Russanov, 1968; Sher, differs from scotti Yakutia can be dated to the Late Middle or the 1971). C. postremus C. in the The former beginning of the Late Pleistocene. Earlier, this shape of the palmation (fig. 4).

well C. has an undivided, point of view was expressed by Russanov species (as as latifrons) Scott's has (1968), and Vangengeim (1977). Sher (1986) flat palmation, while stag-moose a defined the biozone of this broad-fronted moose bi- or tripartite palmation, with its parts often antler beams - in different as the Middle Pleistocene (Mindel Riss, Riss), lying planes. Besides, of shorter and thinner and, likely, the beginning of the Late Pleisto- C. postremus are slightly than C. scotti - in cene (Riss Würm). (table 12). Specimen YIGS1277 is the only presently Description and comparison. Antler fragments known skull of C. postremus. It is noteworthy are strongly mineralized, dark brown or grayish that the most important diagnostic area, the brown in colour. Antler beam is long, but relati-

45 A review of the of Pliocene and Pleistocene 1 systematics moose, part

Fig 13 Skull fragment of Cervalces postremus from Verkhneviliuyskoye outcrop, the Viliuy river (YIGS 1277): a -

- of frontal elevation marked Scale bar view from above, b posterior view (the area is by an arrow). 1 cm

Schedelfragment van Cervalces postremus uit Verkhneviliuyskoye ontsluiting aan de Viliuy rivier (YIGS 1277): a -

b - achteren de de is met gezien van bovenaf, gezien van (het gebied van opbolling aan voorzijde aangegeven een

pijl). Maatstreep 1 cm

basal part of the frontals(fig. 12b), is preserved. Alces (table 5); besides, the labial pillars are The antler beam of this has the which feature for the specimen present, is an obligatory

characteristic size and proportions of the ascription to Cervalces.

“postremus ” form (table 11). Moreover, an exten- Remarks. Sher (1986) and Bubenik (1986) sive distance from the burr to the midpoint of assumed that the form “postremus” is probably the frontal, and the weak frontal protuberance within the an independent species genus confirm the attribution of this clearly moose to Cervalces. Sher (1986) and Lister (1993) argued the Cervalces. The distance from the burr genus that due to the lack of a skull of the “postremus” to the mid-frontal (the fronto-parietal suture) in form, its attribution to Alces or Cervalces is YIGS1277 is which is much specimen 12,5 cm, obscure. The skull fragment YIGS1277clearly more than in Alces (table 2). In fossil and shows that “postremus”, the broad- subfossil this from younger true moose ranges parameter fronted related moose, is not to true moose and with of 9,6 to 10,4 cm, a mean 10,1 cm (n=6); to the Cervalces. This belongs genus can already recent forms show of to a range 8,8 11,1 cm, be concluded on the basis of antler beam with a mean of 9,69 cm (n=16). morphology, because their proportions are The mandible fragments YIGS 4406and YIGS characteristic for the lineage of broad-fronted 4436 are preliminary referred by us to C. moose (table 11). Besides, antler remains of C. postremus, since their molars have large dimen- C. indicate broad postremus a and undivided sions, intermediate between C. latifrons and palmation with long tines (fig. 4b), similar to the

46 CRANIUM 22, 2 - 2005

Table and index of antler beams of the 12 Sizes (in mm) massivity (= 3:1, in %) stag-moose Cervalces scotti and C.

- cf. scotti fromAlaska (coll. AMNH). 1 length; 2 - circumferenceof the burr; 3 - circumferenceabove the burr; 4 -

circumference at midpoint; 5 - horizontal diameter.

massiviteitsindex Maten (in mm) en (= 3:1, in %) van geweistangen van hert-eland Cervalces scotti en C. cf. scotti

uit Alaska (coll. AMNH). 1 - lengte; 2 - omtrek rozenkrans; 3 - omtrek boven rozenkrans; 4 - omtrek halverwege; 5

- dwarsdoorsnede.

Measure- AM 3422426 AM 2768309 AM 5155170 AM 2358130 AM 2766360 AM 2304231 AM 2758313 AM 1825414 AM 2297894 ments

1 290 254 340 305 330 205 220 265 220

2 280 231 300 220 240 170 240 235 185

3 201 189 250 180 210 149 185 220 160

4 185 170 180 158 176 139 180 183 152

5 57,4 55,4 57 49,5 57 43 59 57 46,8

3:1 69,3 74,4 73,5 59 63,6 72,7 84,1 83 72,7

the morphology known for C. latifrons. There is Boeskorov, 2001), and Kamchatka (Vaskovsky,

time a significant gap between C. latifrons; and 1966), the actual distribution area of this moose

C. postremus. Remains of the former species are was supposedly much wider. According to

common in of the and Alekseeva the antlers of Pleisto- deposits Early early (1980), among Middle Pleistocene of Northern Eurasia. The cene moose from the Krasnyi Yar locality (the

latter species is characteristic for the deposits of Ob River region) there are specimens with a

the late Middle Pleistocene of Eastern Siberia beam to 26 This material length up cm. may and the Russian Far East, whereas C. latifrons is indicate C. postremus. Vereshchagin (1967)

already absent at this stratigraphic level described a moose antler fragment from the (Baryshnikov et al., 1981; Sher, 1986). The size of MiddlePleistocene deposits of the Volga River

antler beams of the late broad-fronted moose is region. Similar to C. postremus, this specimen is much smaller than known for C. latifrons, and thin (the diameter above burr is 40,5 cm), and almost no overlap is observed (fig. 15). The its beam has an average length (24 cm). Fossil and antlers of the temporal morphological discontinuity moose “postremus ” type are between C. latifrons and C. postremus clearly known also from the eaerly Late Pleistocene of indicates that latter cannot be a subspecies of Ehringsdorf, Germany (Sher, 1986). the broad-fronted but moose, represents an Cervalces sp. independent species. Material and Mandible Judging from its sites, C. postremus was widely locality. AM 108-5745, distributed from Central Yakutia and the delta Alaska, Engineer Creek (AMNH). of the Lena River the Geological This mandible is referred to in up to Kolyma Lowland, age. the Western Chukotka, and Kamchatka (fig. 5). the AMNH catalogue to Rancholabrean, the

The of in from Middle to Late Pleistocene. The area C. postremus Eastern Siberia was period quite similar to that of C. latifrons. However, the bone is mineralized and has a dark brown colour. younger broad-fronted moose obviously expanded still further to the east, and it is likely Description and comparison. The dimensions of the described bone table that in the late Middle Pleistocene C. postremus are presented in 3.

dispersed from the extreme northeast of Asia Proportions of this jaw (indices of dentition along the Beringian Bridge into North America length, diastema length, height at coronar to give rise to C. scotti. In Eurasia, the late process) reveal that it belongs to Cervalces. At broad-fronted the same time the of of moose, apparently, got extinct in majority parameters the beginning of the Late Pleistocene. This AMNH 108-5745 is lower than in broad-fronted have been influenced exceeds those of Scott's process may by moose of moose, though stag- the to cold moose. this mandible to genus Alces: being more adapted a Probably belongs an could intermediate form between C. and C. climate, they displace C. postremus. latifrons close Although the most reliable finds of C. postremus scotti, to C. postremus. are known from Eastern Siberia (Yakutia) (Vangengeim & Flerov 1965; Russanov, 1968;

47 of A review of the systematics Pliocene and Pleistocene moose, part 1

14 of of - MamontovaGora b Fig Fragments antlers Cervalces postremus. a typical specimen, (after Russanov, 1968);

- YIGS 45, Mamontova Gora; c - YIGS 135, Rossypnoye locality; d, e - YIGS 970 and YIGS1310,

Verkhneviliuyskoye locality; f - YIGS 4758, the Lenariver delta

Geweifragmenten van Cervalces postremus. a - karakteristiek exemplaar, Mamontova Gora (naar Russanov, 1968);

b - YIGS - - YIGS 970 45, Mamontova Gora; c YIGS 135, Rossypnoye vindplaats; d, e en YIGS1310,

f - YIGS delta de Lena Verkhneviliuyskoye vindplaats; 4758, van

Cervalces scotti Lydekker, 1898 - Scott’s stag- Creek; antler beams: AMNH 182-5414, 235-8130 276-8309 moose (fig. llg), 275-8313, (fig. llf), Engineer Creek; AMNH 229-7894and 515-5170 (fig. llh), Cervalces roosevelti Hay, 1913 vicinity of Fairbanks; AMNH 230-4231, Eldo- C. borealis Bensley, 1913 rado Creek; N 276-6360, Banks-at-Fox (fig. lie); Material and locality. Collections of AMNH. A AMNH 342-2426, Cripple Creek. skull (fig. 16a), mandible (fig. 7c), metatarsal, Material of Scott's AMNH 107678, New Jersey, Mount Hermon. Geological age. stag-moose is referred Alaska: skull (fig. 16a) and mandible (fig. 7d), in the AMNH catalogue to the Rancho- the of AMNH 198-5899, Engineer Creek; skull frag- labrean; geological age this material is not determined. All osteal of C. ments: AMNH 101-5090 (fig. 17b) and AMNH exactly remains

scotti are mineralized and in colour from 627-6546, both from Engineer Creek; AMNH range

dark brown to 342-2426, Cripple Creek (fig. 17c); AMNH up yellowish-brown. A similar of mineralization is also found bones 712-1342and AMNH 16250, vicinity of Fair- degree in from the Holocene. banks; mandible AMNH 108-5745, Engineer Early The skeleton from

48 CRANIUM 22, 2 - 2005

Fig 15 Antler fragments of Cervalces postremus. a - private collection, no number, Adycha river, Ulakhan-Sullar; b

- Northern Yakutia. For - YIGS - YIGN, no number, comparison Alces cf. americanus: c 631, Mamontova Gora; d

MM 7653,Kolyma river, Duvanny Yar; e - YIGS 3380, Bolshaya Chukochiya river (after Lazarev & Tomskaya, 1987)

Cervalces - zonder b - Geweifragmenten van postremus. a privé collectie, nummer, Adycha rivier, Ulakhan-Sullar;

YIGN, zonder nummer, Noord-Yakutië. Ter vergelijk Alces cf. americanus: c - YIGS 631, MamontovaGora; d - MM

7653, Kolyma rivier, Duvanny Yar; e - YIGS 3380, Bolshaya Chukochiya rivier (naar Lazarev & Tomskaya, 1987)

New AMNH 107678 found in which come into contact with the Jersey was a 13), upper of the These post-glacial bog (Kurtèn & Anderson, 1980), so processes premaxillary bones, etc. it probably belongs to the beginning of the features indicate that stag-moose and broad- fronted uniform Holocene. The skull with mandible AMNH moose belong to a genus 198-5899is mineralized and also charred: it Cervalces (Azzaroli, 1981). bears traces of burning by fire, most likely Skull AMNH 198-5899was not described yet. caused humans. basic rather similar those by prehistoric According to Its parameters are to recent data people penetrated into Alaska some of a skull of C. scotti from New Jersey (table 13), the 12.000 -13.000 years ago (after Guthrie, 1990a); which undoubtedly belongs to same species the of this be and the existence of Scott's geological age stag-moose can proves stag-moose in determined as the end of the Pleistocene or the Alaska. Specimen AM 198-5899is one of the beginning of the Holocene. Our C. scotti mate- few most informative remains of C. scotti, and rial thus from this allowed with skull AM probably originated period. us, together 107678, to

test the reliability of some features with which Description and comparison. Though the stag-moose can be distinguished from true skeleton of a stag-moose from New Jersey was and add additional moose, to diagnostic investigated by Scott (1885) and Hay (1914), a features. detailed metrical description of its skull and There seems to have been a synchronic mandible was never made. Our data on the existence at the end of the Late Pleistocene and, metrical features of AMNH 107678are given in apparently, in the beginning of the Holocene in tables 3 and 13. The skull of this specimen North America of two representatives of the revealed distinctive features of stag-moose, subfamily Alcinae: Scott's and which stag-moose puts it together with the broad-fronted which resemble each other American moose, moose and distinguishes it from Alces: short in features of skull and significantly many post- rostrum, long nasal bones (which length cranial skeleton. This circumstance considerably exceeds the of the length upper dentition) (table

49 A review of the systematics of Plioceneand Pleistocene moose, part 1

height index of the occiput in broad-fronted

moose is 69,9 %, and in C. scotti 64,8 to 71,3 %,

with of % In an average 68,3 (n=5) (table 1). fossil and modernAlces the occipital part is

higher, with an indexof more than 70%) (table 2).

Moreover, we discovered that the transcornual

width of the skull is more than in the true

In it moose. the former is 11,2 to 12,2 cm, with

of in the latter it is an average 11,54 cm (n=5);

in less than110 mm. Even the largest fossil and Alaska this subfossil Alces americanus from para- between and meter varies 10,2 cm 10,65 cm,

with an of 10,51 cm average (n=6) (table 2).

Cervalces, in contrast to Alces, had a more level

intercornual without the area, pronounced,

cone-like prominence (fig. 17).

16 Sizes of antlerbeams of moose fromthe Fig The diastema of Scott's stag-moose was shorter Pleistocene of Eastern Siberia and the Russian Far East: than in true moose (table 3), thus the index of

- - - Alces A. a Cervalces latifrons; b C. C. postremus; c sp. and. diastema length in the former is close to that in cf. americanus C. latifrons, which is less than 38%. In Alces the

diastema is much because of the eland het longer, general Maten van geweistangen van uit Pleistoceen of the rostral of skull, and here - lengthening part van Oost-Siberië en het verre oosten van Rusland: a

the index is than 38%; moose from Nort- b - C. - Alces A. cf. higher Cervalces latifrons, postremus, c sp. en reach indices americanus heast Siberia and Alaska even

higher than 39% (table 4). broad-fronted For the stag-moose, as well as for of fossil and complicates the diagnostics many of the lower moose, the rather large length of Alcinae this subfossil bones in territory, espe- dental row (index higher than 35%) and the cially so because the rather rare finds consists height of the mandible at the coronar process mostly of neurocraniums, whereas Cervalces and (index higher than 49%) are characteristic (table Alces mainly differ in the nasal bones and the have shorter denti- 3). True moose a relatively rostral part of skull. tion (index usually lower than 35%) and a lower of skull For Cervalces a wide occipital part is mandible (index usually lower than 48%) (table characteristic (after Vislobokova, 1990), together 4). with of the The a small height occiput (fig. 18)

Table 13 Sizes (in mm) of skulls of Cervalces scotti and Alces fromAlaska

Maten(in mm) van schedels van Cervalces scotti en Alces uit Alaska.

Species, coll. Common Condilo-basal Upper toothtooth Cheek bonesbones Rostrum Orbito-facial Nasal bone Height at Rostrum width number, locality length lengthlength row length width length lengthlength length nasion point

Cervalces scotti, Q3-Q4

AMNH107678,AMNH 107678, 552 525266 144 229 175 323323 181 142 8282 New Jersey

AMNH 198-5899,

- Alaska, Engineer 500 - 139 215,3 - -- 155 144

Creek

Alces americanus, Alaska, Q3-Q4

AMNH 106-7527, 6U614 5g4584 152 232 2822g2 3g6386 122 152 g585 Ester Creek

AMNH 368-1426, _- - 159 - 296 409 - - 73 Fish Creek

AMNH 485-4734, 6Qg608 573 152 216216 278 378 104 146 80 Hunter Creek

50 CRANIUM 22, 2 - 2005

Fig 17Skulls of Alcinae from the Late Pleistocene of North America, a - C. scotti, AMNH 107678, New Jersey,

- Mount-Hermon; b - C. scotti, AMNH 198-5899, Alaska, Engineer Creek; c A. americanus, AMNH 106-7527,

Alaska, Ester Creek

Alcinae het Laat Pleistoceen Noord-Amerika, - C. AMNH New Schedels van uit van a scotti, 107678, Jersey,

- - A. AMNH Mount-Hermon; b C. scotti, AMNH 198-5899, Alaska, Engineer Creek; c americanus, 106-7527, Alaska, Ester Creek

Scott's from The premolars to molars ratio in stag-moose, as Beam length in stag-moose varies & The in broad-fronted moose and true moose, varies 30,9 to 41,7 cm (Churcher Pinsof, 1987). from Alaska highly (tables 3 and 4). C, scotti has large tooth length of our antler beams ranges

from 34 with an of 26,99 cm (table 5), and in the mandibles examined by us, 20,5 to cm, average the molars have labial pillars. (n=9) (table 12). The massivity index of the

Antlers. C. scotti has large antlers with a long, antler beam is less than the 100% that is in but relatively thin beam and the original palma- general characteristic for Cervalces, and varies tion divided into two or three parts, with many between 59,0 and 84,1%, with an average of long tines (fig. 4c). A similar antler form is not 72,5% (n=9), which is rather similar to the same found Alcinae. The of C. in C. The among other spread parameter postremus (table 11). circumference of the antler beam of Scott's scotti antlers, apparently, is very large: the stag- moose even the at from 13,9 to 18,5 from New Jersey, which is not stag-moose midpoint ranges has antler of with of and largest specimen, an spread 1,62 m cm, an average 16,92 cm (n=9), thus (Hay, 1914). Such dimensions of antlers testify slightly exceeds the value of this parameter in that C. scotti did not inhabit dense woods. C. postremus.

51 A review of the systematics of Pliocene and Pleistocene moose, part 1

- - b Fig 18 Occipital view of the Alcinae skulls from the North America. C. scotti: a AM 107678, New Jersey,

- AMNH d - AMNH AMNH 198-5899, Alaska, Engineer Creek, c 342-2426, Creeple Creek, 101-5090,Engineer

- - - of Fairbanks. A. Alaska: d AMNH AM Creek, e AMNH no number, vicinity americanus, 106-7527, e 485-9734,

- - 615-1108 f AMNH 508-1489, g AMNH

Achteraanzicht uit C. scotti: - AMNH New b - AMNH van Alcinae schedels Noord-Amerika. a 107678, Jersey,

- - d - AMNH 198-5899, Alaska, Engineer Creek, c AMNH 342-2426, Creeple Creek, 101-5090, Engineer Creek, e

- Fairbanks.A. americanus, Alaska: d - AMNH AMNH f AMNH zonder nummer, omgeving 106-7527, e 485-9734,

- - AMNH 615-1108 AMNH 508-1489, g

till from the Middle C. scotti had body proportions generally similar 33.000 10.000 years ago,

short Wisconsinan to the border betweenPleistocene to those of true moose: a rather neck, long

and Holocene. concomitant fauna body, very long legs and a high body height Dating on for the of the till (Scott, 1913; Hay, 1914; Sher, 1971); the skeleton allows extension existence somewhere the of the Holocene from New Jersey has a withers' height not less in beginning & than 1,81 m). Thus their metapodial length even (Churcher Pinsof, 1987). the for Alces exceeds average parameters (Hay, To be continued 1914) (tables 8 and 9).

North For the Alces and the conclusion, see the Scott's stag-moose remains in America genus next issue of CRANIUM (23,1), scheduled for already originate from the Sangamonian inter- 2006. glacial (Middle Pleistocene) (Kurten & spring

Anderson, 1980). However, the most reliable remains of this species are characteristic only

for the Late Pleistocene, Late Rancholabrean

(Martin & Steadman, 1999). Guthrie (1990b) provided two radiocarbon dates for remains of Cervalces (presumably, C. scotti) from Alaska

+ (vicinity Fairbanks): 32.040 870 - 980 years BP

and + 200 BP (DIC-3090) 25.330 y. (AA-3896).

The dates for C. scotti 11.230 +160 youngest are

BP y. (New Jersey, Colombia; Harington, 1984),

10.950 + 150 BP y. (New York, Middleton;

& and 10.230 + 150 BP Buckley Willis, 1970) y. (Ohio, Ansonia; Meltzer & Mead, 1985). Thus,

the radiocarbon data testify that Scott's stag- moose lived inNorth America at least from

52 CRANIUM 22, 2 - 2005

References

Alexeeva, E.V., 1980. Mammalsof the Pleistocene of Chernyavsky, F.B. & V.I. Domnich, 1989. Moose in

the south-east of the Western Siberia. Moscow: the North-East of Siberia. Moscow: Nauka

Nauka 185 In Russian. 125 In Russian. Publishers, pp. Publishers, pp.

Anonymous, 1987.The Decisions of the Churcher, C.S. & J.D. Pinsof, 1987. Variationin the

interdepartamental stratigraphic session on the antlers of North American Cervalces (Mammalia;

Quarternary system of the East of USSR (Magadan, Cervidae): review of the new and previously 1982): 64-65. Magadan: Publishing houseof recorded specimens. Journal of Vertebrate SVKNIIDVO of the Academy of Sciences of USSR. In Paleontology 4: 373-397. Russian. Danilkin, A.A., 1999. Deer (Cervidae). Moscow: H.E., 1928. Field book of North American GEOS 552 Anthony, Publ., pp. mammals. New York, 674 pp. Erdbrink, D.P., 1954. On one of the oldest known

Azzaroli, A., 1952. L'Alce di Seneze. Paleontografia remains of the common elk, Alces alces L., found Italica 47:133-141. recently in the Netherlands. Geologie en Mijnbouw,

nieuwe serie 16: 301-309. Azzaroli, A., 1981. On the Quaternary and Recent

cervid Libralces. Bolletino 1983. Los' generaAlces, Cervalces, Filonov, K.P., (Moose). Moscow: Lesnaya

dellaSocieta Italiana 71: 48-54. 274 In Russian. Paleontologica promyshlennost Publishers, pp.

Baryshnikov, G.F., V.E. Garutt, I.M. Gromov, Flerov, K.K., 1931. A review of the Elks or Moose A.A.Gureev, I.E. Kuzmina, A.S. Sokolov, P.P. (Alces Gray) of the Old World. Doklady Akademii

Strelkov, A.V. Godina & V.I. Zhegallo, 1981. Nauk SSSR 1: 71-74. In Russian. Catalogue of the Mammalsof the USSR Flerov, K.K., 1934. On geographic distributionand (Pliocene-present). Leningrad: Nauka Publishers, 456 systematics of moose. Doklady Akademii Nauk In Russian. pp. SSSR 2:1-7. In Russian.

Baigusheva, V.S., 1971. Fossil fauna of the Flerov, K.K., 1952. The musk-deer and deer. The Liventsovsk quarry (north-eastern Asov region). fauna of USSR. Mammals. V.l. Issue 2. Moscow, Proceedings of the antropogene of the USSR. Nauka 256 Leningrad: Publishers, pp. In Russian. Leningrad: Nauka Publishers: 5-43. In Russian. Flerov, K.K., 1979. Bison Hamilton Smith. In: V.E. Bobrinskiy, N.A., B.A. Kuznetsov & A.P. Kuzyakin, Sokolov (ed.). Aurochs. Morphology, taxonomy, 1944. A guide to the USSR Mammals. Moscow: evolution, ecology: 4-48. Moscow: Nauka Publishers. Sovetskaya Nauka Publishers, 439 pp. In Russian. In Russian.

Boeskorov, G.G., 1996a. Chromosomal

K.K. & N.C. 1963. Pseudalces - Flerov, Shevyreva, a differentiation on Moose (Alces). Proceedings of the Pleistocene deer from Cis-Caucasia. Paleontological Russian Academy of Sciences 348, 2: 275-278. In Journal 3:130-133.In Russian. Russian.

Frick, Ch., 1937. Horned ruminantsof North Boeskorov, G.G., 1996b. On Taxonomical structure of America. Bulletinof the American Museum of the Genus Alces (Artiodactyla, Cervidae). Vestnik NaturalHistory 69:1- 669. zoologii 6: 70- 78. In Russian.

Geist, V., 1998. The Deer of the World: their Boeskorov, G.G., 1997. Chromosomal differences in evolution, behaviour, and ecology. USA: Stackpole Moose (Alces alces L., Artiodactyla, Mammalia). 421 books, pp. Russian Journal of Genetics 33, 7: 825-828.

Gromov, I.M., A.A. Gureev, G.A.Novikov, Boeskorov, G.G., 1998. Differentiation and the I.I.Sokolov, P.P.Strelkov & K.K.Chapsky, 1963. The problem of Moose (Artiodactyla, Cervidae, Alces) mammalsof the USSR fauna, part 2: 641-2000. systematics. Russian Journal of Zoology 2, 2: 350-362. Moscow, Leningrad: Publishing houseof the Boeskorov, G.G., 2001. The systematics and origin of Academy of Sciences of USSR. In Russian. the modernmoose. Novosibirsk: Nauka, 120 pp. In 1965. A brief of the Gromova, V., survey Quaternary Russian. mammalsof Europe. Moscow: Nauka Publishers, 1999. New data Boyeskorov, G.G., on Moose (Alces, 141 pp. In Russian. Artiodactyla) Systematics. Saugetierkundliche Groves, C.P. & P. Grubb, 1982. Relationships of Mitteilungen 44,1: 3-13. living deer. Biology and management of the Bubenik, A.B., 1986. Taxonomic position of Alcinae Cervidae: 21-59. Washington: Smithsonian Institute

1874 and the of the Alces Press. Jerdon, history genus Gray, 1821. Alces 22:1- 67. Guthrie, R.D., 1990a. Frozen Fauna of the Mammoth

steppe. Chicago and London: The University of 323 Chicago Press, pp.

53 of the of Pliocene and Pleistocene 1 A review systematics moose, part

dates Alaskan 1980. The horses of Guthrie, R.D., 1990b. New on Lazarev, P.A., anthropogene

- Nauka 189 In Quaternary Moose, Cervalces-Alces Archaeological, Yakutia. Moscow: Publishers, pp. Evolutionary, and Ecological Implications. Current Russian. Research in the Pleistocene 7:111-112. Mammalsand Lazarev, P.A. & A.I.Tomskaya, 1987. Northern Haltenorth, Th., 1963. Klassifikation der Saugetiere: biostratigraphy of the Late Cenozoic of of Yakutian Artiodactyla. Handbuch der Zoologie 8,32:1-167. Yakutia. Yakutsk: Publishing House

Scientific Centre, 169 pp. In Russian. land Harington, C.R., 1984.Quaternary marine and mammalsand their paleo-environmental Lister, A., 1993. Evolution of mammoths and moose:

- from northernNorth the Holarctic in implications some examples perspective. Morphological change America. Carnegie Museum of Natural History Quaternary mammalsof North America. Special Publications 8: 511-525. Cambridge: University Press: 178-204.

Hay, O.P., 1914. The Pleistocene mammalsof Iowa. Lydekker, R., 1915. Catalogue of the ungulate Cervalces. Iowa Geological Survey, Annual Report for mammalsin the British Museum (Nat. Hist.). V. IV. London: British 1912: 261-269.Des Moines: Iowa Geological Survey Artiodactyla, fam. Cervidae.

Print:261-269. Museum, 438 pp.

P.S. & D.W. 1999. Heintz, E., 1970. Les Cervidae villafranchiens de Martin, Steadman, Prehistoric Islands and Continents. In: Ross & France et d'Espagne. Memoires du Museum Extinctions on National d'Histoire Naturelle 22:1-303. MacPhee (eds). Extinctions in Near Time: 17-56. New York, Boston, Dordrecht, Moscow: Kluwer Heintz, E. & F. Poplin, 1981. Alces carnutorum Academic/Plenum Publishers. (Laugel, 1862) du pleistocene de Saint-Prest (France). from Alaska. Systematique et evolution des Alcines (Cervidae, Miller, G. S., 1899. A new moose Mammalia). Quartarpalaontologie 4:105-122. Proceedings of the Biological Society of Washington

13: 57 - 59. Heptner, V.G., A.A. Nasimovich & A.G. Bannikov, 1961. Mlekopitayushie Sovetskogo Soyuza Naniev, V.I., 1956. The Caucasian moose, "saerdgyn (Mammals of theUSSR). Artiodactyla and sag" (Alces alces caucasica subsp. nov.). Scientific Perissodactyla. Moscow: Vys'shaya Shkola notes of North. Osetinsky state pedagogical insitute

1004 In Russian. of K.L. 20:151-167. Ordzhonikidze: Publishers, pp. Khetagurov North Osetinsky Publishing house. In Russian. of Deer Johnson, R., 1874. Notice of a new Species sound from the Norfolk Forest-Bed. Ann. Mag. Natur. Hist. Nikolsky, A.A., 1984. The mammal signals in the evolution Moscow: Nauka 4,13: 2. process. Publishers, 199 In Russian. pp. Kahlke, H.D., 1990. On the evolution, distribution The archaic and taxonomy of fossil elk/moose. Nikolsky, P.A., 1996. broad-fronted of the The Quartarpalaontologie 8: 83-106. moose Kolyma lowland (North Yakutia).

theriofauna state in Russia and neighbouring Kahlke, H.D., 1999. The history of the origion, counrties: 247-253. The internationalsession works. evolution and dispersal of the Late Pleistocene Moscow: Publishing house of Rosselkhozacademy. Mammuthus-Coelodontafaunal complex in Eurasia In Russian. (large mammals). USA, Rapid City: Fenske 1906. Etudes L'histoire Companies, 219 pp. Pavlow, M., sur Paleontologique des Ongules. Sankt-Petersburg, 95 1948. Kaplanov, L.G., Tiger, izubr, moose. Moscow, pp. 128 In Russian. pp.

Peterson, R. L., 1955. North American moose. Kotov, A.A., V.G. Vlasov, V.I. Akhchagnyrov, P.P. Toronto: of Toronto 280 University Press, pp. Philippov & V.N. Olkhovnik, 1976. The report of the work results transsections of R. 1974. A review of the life of study on the basic Peterson, L., general Nat. Canadien 101: 9-21. the Cenozoic sediments in a basin of the Adycha history of moose. river aimed for working out a paleontologic Repenning, Ch., 1980. Faunal exchanges between basementof the regional biostratigraphic schemes of Siberia and North America. Canadian Journal of the Verkhoyanie Cenozoic. Yakutsk, 224 pp. In Anthropology 1: 37-44. Russian. Unpublished manuscript. 1983. William Hunter Rolfe, W.D.I., (1718-1783) on Kozhamkulova, B.S.,1974. The broad-fronted moose Irish "elk" and Stubbs's Moose. Archives of Natural (Alces latifrons Johns.) of Zabaikalya. V.II: 89-92. History 11: 263-290. Novosibirsk: Nauka Publishers, Siberian department. In Russian Rouillier, Ch., 1846. Etudes Paleontologiques sur les Environs de Moscou. Jubilaeum semisaecularem Kurten, B. & E. Anderson, 1980. Pleistocene Doctoris Medicinaet Philosophia G. Fischer de mammalsof North America. New York: Columbia Waldheim: 1-14. Moscou. University Press, 442 pp. Russanov, B.S., 1968. Biostratigraphy of the Late

Cenozoic sediments of SouthernYakutia. Moscow:

Nauka Publishers, 459 In Russian. pp.

54 CRANIUM 22, 2 - 2005

fossil Scott, W.B., 1885. Cervalces americanus, a moose Vislobokova, I.A., 1990.Fossil deer of Eurasia.

or elk, from the Quaternary of the New Jersey. Proceedings of the Paleontological Institute Proceedings of the Academy of Natural Science 37: (Moscow) 240:1- 207. Moscow: Nauka Publishers. 181-202. Philadelphia Vislobokova, I., 1996. The Pliocene Seton, E. T., 1910. Life-histories of Northern animals. Podpusk-Lebyazh'e mammalianfaunasand VI Grass-eaters. London: Constable & Co Ltd., 673 assemblage, Western Siberia. Paleontographia Italica

83:1-23. pp.

Sher, A.V., 1971. Pleistocene mammals and Vislobokova, I., E. Dmitrieva & N. Kalmykov, 1995. stratigraphy of the Far Northeast USSR and North Artiodactils from the Late Pliocene of Udunga,

America. Moscow: Nauka Publishers, 310 pp. In Western Trans-Baikal, Russia. Journal of Vertebrate Russian. Paleontology 15:146-159.

and evolution of The of the Sher, A.V., 1986.History moose. Vorontsov, N.N., 1958. significance

Biologiya i ispolzovanie losya: 6 - 35. Moscow. In chromosome sets study for systematics of mammals. Russian. Bulletinof the Moscow Society of Naturalists 63,

2:5-36. In Russian. Soergel, W., 1912. Das Aussterben diluvialer Saugetiere und die Jagd des diluvialenMenschen. Wilson, D.E. & D.A.N. Reeder (eds.), 1992. Mammal Festschrift XLIII Allg. Vers. Deutsches Species of the World: A Taxonomic and Geographic

Anthropologishes Gesellschaft zu Weimar 2:1-81. Reference. Washington: SmithsonianInstitute Press,

2nd edition, 1312 pp. Sokolov, 1.1., 1959. The hoofed mammals. The fauna

of USSR 1, 3:1-639. Mammals. Moscow, Leningrad:

Nauka Publishers. In Russian.

Vangengeim, E.A., 1961. Paleontologic foundation of the Anthropogene stratigraphy of the Northern Asia. Moscow: Nauka 172 Publishers, pp. In Russian.

Vangengeim, E.A., 1977. Paleontologic basement of

antropogene stratigraphy of the NorthernAsia. Moscow: 172 In Russian. pp.

Vangengeim, E.A. & M.V. Sotnikova, 1981. Geology and fauna of the Zasukhino site. Bulletinof the

Commission for Quaternary period study 51:

106-117. In Russian.

Vangengeim, E.A. & K.K. Flerov, 1965. Broad-fronted Siberia. moose (Alces latifrons) in

Bulletin of the Commission for Quaternary period study 30:166-171. In Russian.

Vaskovsky, A.P., 1966. MiddleQuaternary moose on Kamchatka. Kolyma 1:153-154.In Russian.

Vereshchagin, N.K., 1949. On history and systematic of in the position moose (Mammalia, Ungulata) Caucasus. Proceedings of the Russian Academy of Sciences 56, 3: 491-493 (In Russian).

Vereshchagin, N.K., 1957. The mammalremains from the Lower Quaternary sediments of Tamanian peninsula. Proceedings of the Zoological Institute 22: 9-74. Leningrad. In Russian.

Vereshchagin, N.K., 1967. Geological history of the

moose and its using by prehistoric Man. Biology and

hunting on moose: 3- 37. Moscow: Rosselkhozizdat

Publishers. In Russian.

Vislobokova, I. A., 1986. Elks in the Pliocene of the USSR. Quartarpalaontologie 6: 239-242.

Vislobokova, I.A., 1988. Family Cervidae. In: Biostratigraphy of the Late Pliocene-Early

Pleistocene of Tadjikistan: 72-98. Moscow: Nauka

Publishers. In Russian.

55