CRANIUM 23,1 - 2006

A review of the systematics of and , part 2

Gennady+G. Boeskorov

Summary

This is the second ofthe article theevolution and the ofthe Data from the literature paper part on systematics moose. and of the and data from research New onmorphological variability genetics moose new our own are discussed. data

and on broad-fronted moose (; see part 1) true moose (Alces) is submitted. The former and

direct of.Alces In of revision ofthe of the carried (part 1) are no ancestors (part 2). light new data, a systematics moose is

out, and questions on phylogeny are considered.

Samenvatting

Dit artikel is het tweede deel de de evolutie de de eland. van publicatie over en systematiek van Gegevens uit de

variatie de eland onderzoek worden literatuur over morfologische en erfelijkheid van en gegevens van eigen Ook worden de hert-eland Cervalces fossiele eland besproken. nieuwe gegevens van fossiele (deel 1) en van ware

Alces Cervalces Libralces Aan (dit deel) gepresenteerd. en blijken geendirecte voorouders te zijn van Alces. de handvan nieuwe wordt revisieondernomen de de eland worden betreffendehun gegevens een van systematiek van en vragen phylogenie behandeld.

Family Cervidae Gray,1821 sediments of theOb river (Krasny Yar) are attri-

buted to true moose (Alexeeva, 1980). Moose

bones found in morrain on the Kola peninsula Subfamily Alcinae Jerdon, 1874 probably belong to the Middle Pleistocene Genus Alces Gray, 1821 (Gromova, 1965; Vereshagin, 1967). To the late alces 1758. Type species. Linnaeus, Middle Pleistocene (Tasovskoe glaciation) true

Present, in Europe. moose are assigned connection with a

fauna at the Lower beam of periglacial (tundra-steppe) Diagnosis. average length or Tunguska river (Vangengeim, 1977). short (length always less than circumference).

Braincase is rather narrow and than in higher Pavlov (1906) described an adult true moose Cervalces. The facial of the skull portion is long, skull foundat the Kama river in the vicinity of and the rostrum is Nasal strongly elongated. Mysy village under the name “Alces savinus bones their less than that of are short; length is Fisch." This skull to (fig. 19p). may belong a the dental The of row. upper (nasal) processes Khasarian fauna of the Middle Pleistocene the bones do reach the premaxillary usually not (Gromova, 1965), though its stratigraphic posi- nasal bones. The between the bears area tion requires a re-examination (Sher, 1986). The a cone-like The orbits pronounced prominence. skull shows characteristics of Alces: a long almost level with the frontal vault. are on a rostrum and short nasal bones. Its upper

of bones processes premaxillary are widened, as Species composition. At least two species are in modern European moose. The antler beam is recognised: A. alces (Linnaeus, 1758), early Late long, though much shorter than in broad- Pleistocene - present, Western Eurasia; A. ameri- fronted antlers close the moose; are to -like canus (Clinton, 1822), early Late Pleistocene - type with only a few tines. An original feature present, Eastern (?) Eurasia, end of the Late of this moose is a well-expressed tine on the Pleistocene - North (Wisconsinan) present, right beam (fig. 19p). The development of an America.

additional tine on the beam is not characteristic

Alces sp. for modern moose: a similar formation is not

in found modernAlces, nor of moose by Vereshchagin Remains true (Alces) are found in 136 pairs of antlers), nor us deposits from the late Middle Pleistocene (1949; among by

more than 300 of onwards. Finds of Alces in the Netherlands (among pairs antlers). Appa- sp. rently, there was an original early Pleistocene (Erdbrink, 1954) belong to that period. Some Alces for which the of an additional moose remains from the MiddlePleistocene presence

tine on the beam was characteristic. Similar

3 review of the of Pliocene and Pleistocene 2 A systematics moose, part

Fig 19 Antlers of the fossil and subfossil Alces from Western and Eastern Europe: a - Germany, Schussenried, Late

Würm after b - - (Kahlke, 1956, Sher, 1971); specimen at PMM, Germany, Bavaria, Holocene; c AMNH 27772,

d - - k - Denmark, Holocene; specimen at PMM, Austria, Brück, Holocene; e “Alces palmatus”, Romania,

Pleistocene (after Macarovici, 1961); 1, m - Western Ukraine, Pleistocene (after Vereshchagin, 1967); n -

of - North-East Russia, the Kichmenga river, Pleistocene (?) (after Vereshchagin & Russakov, 1979); o “Alces

of Late Pleistocene-Holocene and - “Alces middlestream of fossilis”, near Moscow, vicinity Mytishchi, p savinus”, the Volga river, lateMiddle Pleistocene (?) (both - after Pavlow, 1906).

Geweienvan fossiele en subfossiele Alces uit West- en Oost-Europa: a - Duitsland, Schussenried, Laat Würm

(Kahlke, 1956,naar Sher, 1971); b - stuk uit PMM, Duitsland, Beieren, Holoceen; c - AMNH 27773, Denemarken,

d - uit - - Pleistoceen Holoceen; stuk PMM, Oostenrijk, Brück, Holoceen; e k “Alces palmatus”, Roemenië, (naar

Macarovici, 1961); 1, m - West Oekraïne, Pleistoceen (naar Vereshchagin, 1967); n - Noordoost Rusland, Kichmen-

- “Alces ga rivier, Pleistoceen (?) (naar Vereshchagin & Russakov, 1979); o fossilis”, omgeving van Moskou, nabij

Laat - “Alces de Mytishchi, Pleistoceen-Holoceen en p savinus”, middenstroom van Wolga, laat MiddenPleisto- ceen (?) (beiden naar Pavlow, 1906).

4 CRANIUM 23,1 - 2006

Antlers of the fossil - Fig 20 and subfossil Alces: a specimen at GM RAS, near Moscow, vicinity of Mytishchi, Ho-

- locene; b, c - GM RAS, Eastern Europe, Holocene (?); d ZM MSU, vicinity of Moscow (?), Pleistocene-Holocene;

e - Stavropol district, near Caucasus region, Holocene (after Vereshchagin, 1949); A. alces cf. caucasicus: f - lower

stream of the Don Middle - Krasnodar the - af- river, Sarkel fortress, Ages, g district, Urup river, Holocene (both

i - Western Pleistocene ter Vereshchagin, 1959); h, Alces sp., , Krasny Yar, (after Alexeeva, 1980).

Geweien fossiele Alces: - stuk in GM van en subfossiele a RAS, omgeving Moskou, nabij Mytishchi, Holoceen; b, c

- GM RAS, Oost-Europa, Holoceen (?); d - ZM MSU, nabij Moskou (?), Pleistoceen-Holoceen; e - Stavropol dis-

de f - trict, nabij Kaukasus, Holoceen (naar Vereshchagin, 1949); A. alces cf. caucasicus: benedenstroom van de

Sarkel - Don, kasteel, Middeleeuwen, g district Krasnodar, Urup rivier, Holoceen (beiden naar Vereshchagin,

i - Alces West Pleistoceen 1959); h, sp., Siberië, Krasny Yar, (naar Alexeeva, 1980).

5 review of the of Pliocene and Pleistocene 2 A systematics moose, part

beams of true found The antler moose were by process among moose. wide, bi-partite Alexeeva (1980) in the Pleistocene sediments of palmation and elongated antler beam is charac-

Western Siberia (fig. 20h-i). Vereshchagin (1959; teristic for the East-Siberian moose of the Late

1967) described the skull of fossil or subfossil Pleistocene. Most probably, the American moose from Cis-Caucasia with rudimental brow moose evolved in Eastern Siberia. tines (fig. 20e). Pleistocene Alces had, on Alces cf. alces and A. alces L., 1758 - moose of average, longer antler beams than modern the European type and the European moose moose (Vereshchagin, 1967). Cervus (Alces) fossilis H. V. Meyer, 1832. Eurasia until True moose were widespread in Alces palmatus Gray, 1843. the of Late Pleistocene beginning (see above). Alces leptocephalus Push, 1846. Probably, during this period their differentia- Material and locality. Separate antlers and frag- tion into two forms, the European and the ments: PMM 1955, Western Germany, Baden- East-Siberian, began. The latter later gave rise to Wurtemberg, Markt-Schwaben; PMM 1950/92, difficult to the North American moose. It is say Bavaria,. Taufkirchen (fig. 19b); PMM 1974/253, where this took In Late exactly process place. Bavaria, Kiinzing; PMM specimen without Pleistocene deposits, true moose remains are number, Austria, Bruk, Markt Grating, (fig. from the found Western Europe up to Russian 19d); AM 27772, Denmark, AMNH (fig. 19c); Far East and Japan (after Kahlke, 1999). Many GM 1823 and GM 1824, Eastern Europe (?), GM finds testify that they were distributed more RAS (fig. 20a, c). Skull fragments with antler widely than at present (see fig. 6 of part 1). parts: ZM 1411, vicinity of Mytishchi Sity (fig.

antler for ZM without of A high variation in shape was typical 20a); specimen number, vicinity Pleistocene Alces (especially in western regions). Moscow (?), ZMMSU (fig. 20d); GM 1829 (?), of the Nerekhta Four antler types are recognized: shovel-shaped vicinity Kostroma Sity, river antlers with undivided palmation, and GM specimen without number, Tiraspol shovel-shaped antlers with bi-partite palmation, (?)• deer-like antlers, and antlers with an additional Geological age. The beam of the left antler the beam. this undoub- tine on In our opinion, PMM 1974/253 is found in the Pleistocene tedly testifies the existence of a differentiation

Table 14 Sizes of fossil and subfossil Alces antlers in of the (in mm); case multiple specimens, averages are given.

* ** Notes: - after specimens from Paleontological Museum of Munich (Germany) and AMNH (USA); - after

specimens from Zoological Museum MSU and Geological Museum RAS, Moscow.

Maten het is in van fossiele en subfossiele elandgeweien (in mm); gemiddelde gegeven het geval van meerdere

* geweien. Opmerkingen: - naar geweien in het Paleontologisch Museumvan München (Duitsland) en AMNH

**

- in Museum MSU (USA); naar geweien het Zoologische en het Geologische Museum RAS, Moskou.

A. cf. america- AlcesAlces alces “Alces"Alces savinus”savinus" A. americanus

nus

North-East

West Germany „ Asia, 0 . Germany . ' Verkhoy- Parameter 3 The Ob villa- Parameter , ~ East Moscow dis- The Ob river, villa- , Europe, river, , . Chersky A1 , and ansk the 3 Denmark, ~. Alaska, Q3-Q4 Denmark, ™ ™ ~, Mnts, A Alaska, Q3-Q4

Q3Q3-Q4" Q4 trict, Q3-QQ3-Q44 Q3-Q4 e Q4 Sge,' Q4, Sartang river,

Q3(?)Q3 (?)

n=4*n=4* n=5**n=5** PINPINRASRAS ZIN 155-895 VMN CMN AMNH

Antler spread ca.1000ca.1000 ca.ca. 1000 ca. 1700 1640 1430 1200 1200,1340

Palmationti° n right-400;left-right - 400; left - 2175217.5 191 caca. 400 right .- 360 300 337.3 (n=ll) width ' 420

Maximal13' 3ntlerantler

left " leftleft-" 9143914.3 902902 ca.ca 1200 Ieft -12601260 1040 1050 1260 ((n=4)n=4) length '

- left - - leftleft -- left - - leftleft - Circum-ference _ . , 220; 210; -190; 205; . „ right right right right 154.8 169Q 183.4 Xa43 (n(n=9)V ofthe beam j 215 214 W0190 200

- left - left -- left - leftleft - 225;225; -120; -150; . , ... r .. right right right right -155; , q, i qo 0 no & Beam length 98.8 129129 128.6 (n=ll) 22155 114 140 150

Number of ti- ?7.8 6.2 12-14 15 14 10 10.8 (n=9)v ' nesnes per antler

6 CRANIUM 23,1 - 2006

Fig 21 Antler shape in European moose: a - from a specimen at KM, Norway; b- from a specimen at MK, Sweden;

d - - Estonia f - c, Western Siberia, Vasyuganie, Demyanka river (after Kaplanov, 1948); e (after Buturlin, 1934);

from a specimen ZM 51087, ZM MSU, Tverskaya district

Geweivorm inEuropese eland: a - naar een stuk inKM, Noorwegen; b- naar een stuk inKM, Zweden; c, d -

West Siberië, Vasyuganie, Demyanka rivier (naar Kaplanov, 1948); e - Estonië (naar Buturlin, 1934); f - ZM 51087 in MSU, district Tverskaya

gravel sediments at a depth of five to ten general, the Late Pleistocene andHolocene The mineralization of the from and had metres. strong speci- moose Western Eastern Europe mens from ZMMSU and GM RAS testifies that antlers with a wide, undivided palmation and they belong to the Late Pleistocene or Early long tines (figs. 19, 20), which is also characte-

Holocene period. AM 27772 and ZM 1411 are ristic for modern A. alces (fig. 21a, c). Only some

in found peat-bogs and are probably of Holo- antlers have a palmation with the tendency to antlers from divided cene age. The other Germany and become into two parts (fig. 19d, j-1).

Austria belong to the Holocene. Futhermore, antlers close to the deer-like type

occur (fig. 19m, o-p; fig. 20e-f), which is also Description and comparison. The antler frag- characteristic for European moose. mentPMM 1974/253 from Bavaria has a rather long (13.5 cm) and massive beam (circumfe- Alces cf. americanus - moose close to the Amer- rence in the middle is 19 cm, circumference ican moose above the burr is 21 The index cm). massivity " Material and locality. Alces savinus”Fischer, (circumference by length) of the antler beam of skull with antlers, without number (?), Moscow 110% proves that this specimen undoubtedly district, PIN RAS (fig. 22a); skull fragment with belongs to Alces. The beam of PMM 1974/253 is antlers ZIN 155-1895, the Ob river, ZIN RAS longer than in modern European moose. PMM (fig. 22b). Mandible fragments, Yakutia: YIGS 1974/253 has a long antler as the brow tine is 4411, the Adycha river; YIGS 4759, the Lena has wide 28.5 cm. Furthermore, it a and, appa- river delta. rently, undivided palmation. The size of the of skulls from the Late Pleistocene The skull of "Savin's moose" occipital part Geological age.

was found in a theRuta river in the - Holocene of Eastern Europe is characteristic peat bog at for Moscow district miles from and Alces (see table 2 of part 1), and their antler (sixty Moscow),

most is from the Holocene sizes are close to those of modernA. alces (table likely period.

14), however, antler beams of the former are Vereshchagin (1967) paid attention to the strong mineralization of this and slightly longer. The parameters of antlers and specimen proposed a Late Pleistocene for the "Savin's moose". beams of Holocene moose from Western age

Europe practically do not differ from those of The label on ZIN 155-1895 tells that it was

found at the Ob The of this exhibit modern European moose (tables 14 and 15). In river. age is

7 A review of the systematics of Pliocene and Pleistocene moose, part 2

15 Sizes in true antler be- Table (in mm; mean case of multiple specimens) and massivity index (in %) of moose

1 - 2 - 3 - circumference above the 4 - 5 - ams. length; circumference of the burr; burr; circumference at midpoint;

horizontal diameter, n.n. = no number.

het meerdere Maten (in mm; gemiddelde in geval van stukken) en massiviteitsindex (in %) van geweistokken van

- - - - eland. - 2 omtrek de 3 omtrek de 4 5 ware 1 lengte; van rozenkrans; boven rozenkrans; omtrek halverwege;

= dwarsdoorsnede, n.n. geen nummer.

"Alces“Alces savi- AlcesAlces alces A. cf. americanusamericanus nus”nus"

Kolyma, DuDu- Bolshaya SizesSizes and in- Germany and Eastern Euro- Moscow dis- Ob river, 03-Q3- Aldan river, Mamontova vanny Yar, Chukarh.yaChukochiya dex Denmark, Q4 pe, Q3-Q4 trict,trict, Q3-Q4 Q4 Gora, Q3 Q3 river, Q3

PMM, ZMMSU, YIGS, MM, PIN RAS, n.n. ZIN155-1895 YIGS631YIGS 631 YIGS778YIGS 778 YYIGSIGS33803380 A AMNH, n=4 GM, n=5 n=5

11 98.8 129 225 JZO120 102 _9595 J105105 90

2 239 233 - 285 270 250 225 250 _- _285 _270 _250

3 173 176.8 250 230 _225225 210 [ 167 J180180

4 154.8 169 J220220 210 JB7187 _180180 152.6152.6 165

48.3 - 57 5 4949.2.2 - 56 47.8 48 _5 j483 _- _57

3:1 207.7207.7 130.8 111.1 191.7 220.6 221.1 172 200

unknown. based rather 15.5 Antler of this However, upon its row cm. sizes moose are

the Museum in tables 14 and 15. strong mineralization, Zoological given Extremely interesting almost certainly will admit it as a fossil. is the fact that although this specimen origi-

Probably this exhibit originates from the Late nated from Eastern Europe, it seems to be Pleistocene the of the Holocene. related the of Northeast or beginning to modern moose

Mandibles are found together with bone Siberia and Alaska by several features. remains of characteristic representatives of the Primarily, these are its craniometric parameters.

Yakutian Mammoth fauna: Mammuthus has antlers, a primige- Additionaly, it very large very nius, Equus lenensis, Bison priscus, Rangifer wide shovel (palmation) with the tendency to

tarandus. Based upon this circumstance, and become divided into two parts. It posesses also the rather mineralization of and antler upon strong many tines very long, massive

the YIGS 4411 and the be beams. the 4759, specimens can The upper processes of premaxillary attributed to the Late Pleistocene. bones, especially the right one, are

wedge-shaped narrowed (see fig. 4e of part I). Description and comparison. The "Savin's This is characteristic for American moose moose" is a remarkable exhibit, now in the exhi- ("American" morphotype, after Boeskorov, bition of the Paleontological Museum of PIN 1998) and very seldom occurs in European RAS. It was originally described as “Cervus of the moose. Besides, in this specimen one () savinus” (Fisher von Waldheim, right shovel tines is directed downwards, which 1834); later as “Alces savinus” (Rouillier, 1846). modern is found in A. among moose only a. Though the "Savin's moose" was already gigas (Bubenik, 1986; our data).The combination well-known and studied (Vereshchagin, 1967; of such an amountof typically American moose Vereshchagin & Russakov, 1979), we re-studied features indicate that 'Savin's moose' must be a it, and found some interesting results. representative of the American group.

Skull and antlers of the "Savin's moose" are The second specimen is equally interesting, and very large. In size they are only comparable to until now undescribed: a skull with antlers on the modern which lives north- largest moose, in exhibit in the Zoological museum ZIN RAS (fig. east Siberia and Alaska and belongs to the 22b). The sizes of the preserved posterior part of subspecies A. a. gigas. Condilobasal length of the skull the width are large: greatest occipital the "Savin's moose" skull was not less than 63 is 17.2 cm. The occipital heights from the lower width in cheek 24 skull cm, bones near cm, of and upper edges the foramen magnum are height in the nasion point 16 cm, length of the 12.8 and 9.7 The width of cm, respectively. nasal bones of 13.6 cm, length the upper tooth occipital condyles 10.1 cm. the antlers of this

8 CRANIUM 23,1 - 2006

and The ular Ob The antlers specimen are very large (tables 14 15). the basin of the river. corre-

majority of parameters of these antlers (spread, spond to the largest antlers of moose from length of antler, width of palmation, circumfer- northeast Siberia and Alaska. The antler shape

ence of beam) is much larger thanthose of the of ZIN 155-1895with the characteristic length-

modern that in ened shovel with the to become European moose occupy partic- tendency

divided into two parts is also typical for

modern amphi-Beringian populations of moose.

antlers of Except for "Savin's moose", some

Pleistocene moose from Western Ukraine and

Middle Volga region are extremely similar to

antlers of modernKolyma and (Vereshchagin, 1967; Vereshchagin & Russakov,

1979). This has led to the conclusion that "...in

the Pleistocene the geographical variability of

other than moose populations was nowadays"

(Vereshchagin & Russakov, 1979: 40). In my

view, however, it seems more likely that during

the Late Pleistocene, the American moose had a

much wider distribution in Eurasia thantoday.

As it is more adapted to a cold climate than the

it have Western European moose, may occupied Siberia and of even parts Eastern Europe. Mandibles YIGS 4411 and 4759 have similar

basic sizes as modernAlces (jaw height,

premolar length, sizes of most teeth; see tables 4

and 6 of part 1), but, at the same time, have

some original features as well. A very large molar length is characteristic for both these

specimens: for YIGS 4411 it is a large length of

the teeth row, and for YIGS 4759 it is the large sizes of M3, similar to Cervalces (see table 3 of

part 1). Among the true moose only fossil,

subfossil and modern A. americanus from

Alaska and the extreme northeast of have more

or less similar sized teeth (see tables 4 and 6 of

part 1), which doubtlessly correlates with their

skulls. very large

Alces americanus Clinton, 1822 - American 22 Antlers of the fossil and subfossil rela- Fig moose moose

tive to A. americanus: a - “Alces savinus”, specimen Material and Skull without numer at PIN RAS, Moscow district, Pleisto- locality. North-east Asia.

cene-Holocene; b - ZIN 155-1895 (ZIN RAS), Ob fragments with antlers: VMN, without number,

river, Pleistocene-Holocene; c - specimen without found in 1996 in Verkhoyansk mountains at the number at VMN, Verkhoyanie, Sartang river, Late Sartang river (fig. 22c); without number, found

Pleistocene (?); d - specimen withoutnumber at in the 1990ties at Chersky village, ChMN (fig. CMN, lower stream of Kolyma river, Chersky vil- 22d). Occipital parts of skulls: YIGS 4291, lage, Holocene. Chukotka, the Malyi Anuyi river, Krassivoye

locality; YIGS two specimens without numbers, Geweien van fossiele en subfossiele eland northern Yakutia MM 8121 and MM two

- (?); vergeleken met A. americanus: a “Alces savinus”, without numbers, Yar stuk zonder nummer in PIN RAS, district Moskou, specimens Duvanny MM 7099 and Pleistoceen-Holoceen; b - ZIN 155-1895 (ZIN RAS), locality. Mandibles: MM, one

de - stuk zonder Ob, Pleistoceen-Holoceen; c specimen without number, the Kolyma river.

in rivier Laat nummer VMN, Verkhoyanie, Sartang, MAE, three mandible fragments, without

Pleistoceen (?); d - stuk zondernummer in CMN, numbers, southwestern Yakutia, the Oleykma de benedenloop van Kolyma, Chersky dorp, river basin, UlakhanSegelennyakh, Late Holo- Holoceen.

9 A review of the systematics of Plioceneand Pleistocene moose, part 2

of the described cene human settlement. Antler fragments: YIGS years ago). The remainder

of and YIGS the from Yakutia have no 631 (see fig. 15c part 1) 788, moose remains geological Aldan river, Mamontova Gora locality; YIGS dating but their mineralisation allows for an

3208, 6395 and one specimen without number, attribution to the Late Pleistocene - beginning of

MM 7313 and MM 7754 (see fig. 15d of part 1), Holocene. The material of the true moose from

Duvanny Yar; YIGS 3380, the Bolshaya the AMNH belongs to the Rancholabrean,

Chukochiya river (see fig. 15e of part 1), YIGS according to the museum catalogue. Bone without northern remains of Alaskan mineralized and one specimen number, moose are

one from dark brown, Yakutia (?); MM specimen without number, their colour ranges light to

the Batamai river; IRMN 6645 and one and can be attributed to the final Late Pleisto- specimen without number, Irkutsk district, near-Baikal region.

Alaska. Skulls: AMNH 106-7527, Ester Creek

(see figs. 17c and 18f of part 1); AMNH

368-1426, Fish Creek; AMNH 485-4734, Hunter

of Skull Creek (see fig. 18g part 1). fragments: AMNH 327-5020, 383-2037, 508-1489 (see fig.

18h of part 1), 615-1107and 615-1108 (see fig. of Fairbanks. Antler 18i of part 1), vicinity frag- ments: AMNH 252-1939, Eldorado Creek (see fig. 12a of part 1), AMNH 4772, 5269, 5291,

252-4442, 276-5295, 292-5273, 292-5293,

292-9143, 348-8487, 420-1360, vicinity of Fair- banks (fig. 23). Mandibles: AMNH 616, of 276-5283 (see fig. 8a part 1), 368-1426, 485-4734, 583-2162, 583-2167, 584-2161, vicinity of Fairbanks. Metatarsals: AMNH 34586,

368-1426, 368-1426/1, 586-2310, vicinity of Fair- banks. Mummified corpse AMNH 274-4002, Little Eldorado Creek (fig. 24).

The skull with antlers Geological age. fragment from the Sartang river obviously belongs to the

Late Pleistocene, because it was found on a

23 of the fossil and subfossil antlers of depth of 2.5 m together with bones of Fig Fragments americanus from Alaska late The A. (coll. AMNH) Mammuthus primigenius of a type. antlers from found Chersky village were at a Fragmenten van fossiele en subfossiele geweien van depth of two metres, so are attributed to the A. americanus uit Alaska (coll. AMNH) Holocene. YIGS 4291, YIGS 3380, MM 7754 and

MM 8121 all originated from the Edoma forma-

tion (Icy Complex) sediments of the Late cene - beginning of Holocene. The moose with is dated 33,000 + Pleistocene and they were found together mummy (AMNH 274-4002) bones of the characteristic representatives of 750 y. BP (Guthrie, 1990b).

this period: M. primigenius of the late type, Coelo- Description and comparison. The measure- donta antiquitatis, Equus lenensis, Bison priscus cf. ments of the fossil and subfossil remains of occidentalis, Rangifer tarandus. The radiocarbon moose skulls from the northeast Asia fall within of MM 7754 from the Duvanny age specimen the the variation of modern moose living in Yar locality (see fig. 15d of part 1) is 12,250 + 70 of Yakutia, but are closer to the territory para- y. BP (GIN -11025). YIGS 631 and YIGS 788 meters of the largest living subspecies: A. a. from the Mamontova Gora locality were gigas. Fossil and subfossil moose from Alaska described by Russanov (1968), and attributed to are outstanding by their greatest sizes (see table the Late Pleistocene, because they were found 2 of part 1). The mandible sizes also testify that in the Late Pleistocene loess-like sediments. The fossil and subfossil Alces from Yakutia belong to mandibles from the Oleykma river basin were the large form. However, Alaskan moose found in the settlement UlakhanSegelennyakh them in sizes table 4 of For surpass (see part 1). layers of the Eneolithic period (1,500 -1,000

10 CRANIUM 23,1 - 2006

24 Late Pleistocene A. americanus found in 1940 Little Creek. Fig mummy (AMNH 274-4002) on Alaska, Eldorado

Laat-PleistoceneA. americanus mummie (AMNH 274-4002) gevonden in 1940 in Alaska, Little Eldorado Creek.

Late Pleistocene and Holocene A. americanus the the Late Pleistocene and Holocene A. americanus

lengthening of molars is characteristic. The sizes represents only beams with the beginning of a remarkable that of teeth of the Yakutian moose fall within the shovel-shape expansion. It is and of them had shovel. The variation of modern A. a. pfizenmayeri A. a. many a bi-partite sizes gigas; the sizes of teeth of Alaskan moose of these beams are given in tables 14 and 15, that the rather and surpass the parameters of the latters (see table 6 and it is obvious long of and table of this antler trunk is characteristic these part 1 16 part). massive for moose. The massivity index of the beam always The mummified of the corpse young moose exceeds 100%. The Late Pleistocene and Holo- from the Little Eldorado Creek (fig. 24) was cene moose from Eastern Siberia and Alaska are Cervalces earlier attributed to latifrons or stag- closest to modern A. a. gigas by their beam Cervalces moose sp. (Guthrie, 1990a, 1990b). length. As Bubenik noted, the morphotype of Nevertheless, our research has shown that this the latter moose (large body and skull, long specimen has characteristics of Alces : a cone-like of antlers with rostrum, huge spread many prominence between the antlers and lengthened tines) developed under conditions of thin nort- hoofs. Absolute of this find + 750 age (33,000 y. hern and taiga-tundra of the northeast BP) is evidence for the penetration of true Siberia and Alaska. In the snow-free period, moose into Alaska prior to the Wisconsinan these moose prefer semi-open to open glaciation. landscapes. Their regular penetrations into the

in The best preserved antlers from Sartang and tundra zone are noted, and Alaska the moose

Chersky (fig. 22c-d) and Alaska (fig. 23) have is even called "tundra-moose" (Bubenik, 1986). the form and this of close the clearly expressed (long wide, Apparently, type moose is to shovel with and Late Pleistocene which of the bi-partite many tines) large moose, was part for modern Mammoth fauna of sizes (table 14), characteristic A. a. in a significant part Nort-

gigas from the Northeastof Siberia and Alaska. hern Eurasia. During the Late Pleistocene, it

The remainder of described antler remains of

11 A review of the systematics of Pliocene and Pleistocene moose, part 2

Alaska the Land last of the Cervalces. Presu- migrated to through Bering representative genus Bridge. mably it became extinct at the beginning of the

Late Pleistocene. Alces from the Pleistocene of Eastern sp. Siberia,

and also fossil and subfossil A. americanus from The salient features of the broad-fronted moose

have which the traced in this evolutio- Alaska very large metatarsals, are (see diagnosis) are

from Pliocene to the end in size closest to those of modern moose from nary lineage the Late

of Siberia table of of the Pleistocene. In view, until now there the Northeast (see 9 part 1). my

is no convincing evidence in favor of a direct

phyletic link between Cervalces and true moose Alces. The latter differ significantly from broad- Conclusion above fronted moose and stag-moose in the Analysis of paleontological data indicates that mentioned features (see the diagnosis). The the evolutionary lineage of the broad-fronted, evolutionary tree of moose is shown in figure known from the long-beamed moose, beginning 25. Most likely the evolutionary lineage of true of the Late Pliocene of Eurasia, apparently got moose developed in parallel with that of extinct in Eurasia the of the Late by beginning broad-fronted the moose, starting as early as Pleistocene, and in North America by the begin- end of the Late Pliocene or the beginning of the ning of the Holocene. Pleistocene. However, remains of a real

ancestral form of Alces have not been found Libralces the earliest known yet. gallicus, representa-

tive of this lineage was replaced by Cervalces Paleontological data to the present time testify of latifrons in the Early Pleistocene. Remains an that a fully-developed true moose already intermediate form the features of L. combining existed by the late Middle Pleistocene. Their

and C. are found in the gallicus latifrons Kolyma evolution took place in the central and northern lowlands 1971; 1986; 1996). (Sher, Nikolsky, altitudes of Eurasia, which rise to a gave broad-fronted with massive Large moose long number of adaptations and features of Alces: antler beams the Middle disappeared during large body sizes, characteristic for deer of nort- Pleistocene of Eurasia. In this period a new, hern altitudes, features of a body structure with smaller form of Cervalces appeared. On average, high withers, lengthened metapodials, enlarged it had a long and rather thin antler beam, and basic area of the hoofs, promoting locomotion was described as the Alces separate subspecies in hillocky bogs, wind-fallen wood and deep latifrons postremus (Vangengeim & Flerov, 1965). snow. Thelarge weight of the nasal cartilages moved to the Cervalcus Later, it was genus and the strongly developed upper lip serve as The and (Sher, 1986). chronological morpholo- but an adaptation to preliminary heating of air, gical hiatus between C. latifrons and the also to grasp foliage from branches and marsh form allow latter postremus- us to consider the as plants (Flerov, 1952; Vereshchagin, 1967). the separate species Cervalces postremus. This Apparently, already at the beginning of the Late species was widely distributed in Eastern radiation Alces took Siberia. Pleistocene, a among place. This is testified by finds of Alces antler remains After the dispersal into North America, it appa- with the various forms peculiar to modern rise to the American rently stag-moose, with with addi- gave moose together unique finds an . In Eurasia C. postremus was the

Table of of modern 16 Sizes (in mm; mean values) lower jaw teeth the moose (genus Alces).

Maten (in mm; gemiddelden) van de onderkaakskiezen van de huidige elk (geslacht Alces)

P2 P P M M P2 P33 P44 MlMI M22 M33 Species and subspecies length width length width length width length width length width length width

AlcesAlces alces dices,alces,Tambov district (Ni- - - 22.5 15.815.8 25.6 17.1 - - 26.5 20 - - kolsky, 1996)

A. dmericdnusamericanus pfizenmayeri,pfizetimdyeri, Central 18 13.413.4 24.2 16.516.5 28.3 18.7 28.4 19.7 29.4 20.5 40 2020 and Southern Yakutia, n= 10

A. d.a. gigas,gigds, Extreme North-East Asia, 19.5 14.214.2 24.5 17 28.7 19.7 28.2 20.6 30.6 22.5 40.5 21.521.5 n=n= 10

12 CRANIUM 23,1 - 2006

Fig 34 Evolutionary lineage of Alcinae / Evolutielijn van de Alcinae.

13 review of the of Plioceneand Pleistocene 2 A systematics moose, part

tional tine on a beam. The forms from which testify that this occupied Eastern Siberia,

later the European-West-Siberian and the East- including its northern part at least from the

Siberian-American moose originated, probably Karginian interglacial of the Late Pleistocene of stood apart at that time. The level of differentia- (more than 40,000 years ago). But remains tion between modern European and American Alces found in the late Middle Pleistocene of the

the basis of Lower river moose on molecular-genetic Tunguska (Vangengeim, 1977)

markers of DNA has allowed to estimate the allows to assume that true moose inhabited the

time of their divergence at more than 100,000 larger part of Eastern Siberia at least from the

& Anderson, or at 75,000 to of the Late Pleistocene. years (Mikko 1995) beginning et 150,000 years (Udina al., 2002). Thus, the Earlier, it was supposed that Alces probably divergence of Alces into two species apparently entered North America from Asia at the begin- occured at the end of the Middle - beginning of of the about ning Wisconsinan, 100,000 years the Late Pleistocene. ago (Hibbard, 1958) or even before the On the basis of palaeontological finds, it is diffi- Sangamonian interglacial (less than 250,000 and cult to which of is the most than say species moose more 100,000 years ago) (Guthrie, 1966). ancient. The presumably earliest remains of late But until now no radiocarbon dated Alces

Middle Pleistocene true moose are found from remains have been found in North America that

Western to Eastern Siberia. At the older than the of Holocene Europe up are beginning

time data that the East-Si- + 70 BP and + 55 same genetic testify (Alaska 8,740 y. 5,380 y. BP,

berian - most after Manitoba and American moose likely are more Guthrie, 1990b; 7,848 y. BP ancient than The + 40 after Brownlee et European moose. karyotype 5,950 y. BP, al, 2002). with 2n=70 is considered as ancestral for the Guthrie (1990b) suggests that Alces penetrated family Cervidae. It is therefore most probable into North America quite recently: between the

that the similar chromosomal set of A. ameri- end of the last and the interglacial (24,000 y. BP)

canus is the archaic set. It was transformed into end of the maximum of the glaciation (14,000 y.

the 2n=68 chromosomal karyotype of A. alces as BP) or even at the beginning of the Holocene

result of Robertsonian fusions & Based the lack of varia- a (Groves (Guthrie, 1995). upon

Grubb, 1982; Grafodatsky et al., 1990). In tion in mitochondrial DNA restriction frag-

Siberia and the the North be Eastern Far East largest ments in American moose, it can variety of haplotypes of mitochondrial DNA is concluded that the population must be rela- revealed. This indicate that this might region tively young and moose entered North America was the closest to the place of origin and evolu- from northeast Asia during the late tion of modern Alces (Hundertmark et al., 2002; Wisconsinan (Cronin, 1992). Udina et al., 2002). In my view, the stronger At the same time there is a fossil yearling bull differentiation of the American moose (five or from Little Eldorado Alaska mummy Creek, six subspecies in Eurasia and North America) (AMNH 274-4002), which is radiocarbon dated than of theEuropean moose also testifies the BP This 33,000 y. (after Guthrie, 1990b). of the former. more ancient origin mummy probably belongs to

East-Siberian moose more a cold Cervalces 1990a, is adapted to or to stag-moose sp. (Guthrie, climate than the former but European moose; was 1990b), more likely to true moose, Alces

distributed the Mammoth fauna This makes it that widely among (Boeskorov, 2001). possible of west-ward Siberia and inhabited Alaska Eurasia, to Western true moose more than thirty even Eastern In the Late Pleistocene, thousand However, the time when Europe. years ago. of this Alces into North America is unclear populations moose were not numerous, penetrated as can be derived from the rare finds of Alces until now. We do known that it must have bone remains at Mammoth fauna sites and at taken place before the end of the Late Pleisto-

Paleolithic settlements. This is the the of because the explained by cene or beginning Holocene, fact that differentiation of open, steppe-like landscapes were not modern North-American

them. Remains of modern three four could suitable for moose moose into or subspecies not found in situ Aldan Mamontova Gora have taken in ten thousand (the river, place years only. locality; after Russanov, 1968), and also radio- For example, the two amphi-Beringian popula- carbon dating (Taimyr Peninsula; after tions of A. a. gigas still did not reach a Sulerzhitsky & Romanenko, 1997) and findings subspecific level of differentiation although at Late Paleolithic sites (Vangengeim, 1977; they were morphologically isolated from each

Lazarev et other for ten thousand Mochanov, 1977; al., 1998) evidently years (Boeskorov, 2001).

14 CRANIUM 23,1 - 2006

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16