CRANIUM 20, 2 - 2003

of the modern and review of its The genetics a taxonomy

Gennady+G. Boeskorov

Summary

results of and of the New data from research The investigations on genetics systematics moose are presented. our own and of the modern and datafrom the literatureon morphological variability, genetics, ecological ethological features discussed. Numerous data that differentiation modernAlces has reached level and moose are testify among aspecific Siberia and they divided into two species: European moose Alces alces L. inhabiting Europe, Ural, and Western

A. americanus Clinton North Eastern and the Far East. These American moose inhabiting America, Siberia, species

has has well differ in chromosome numbers (European moose 2n=68, American moose 2n=70), as as in some

features. of data of the ofthe carried and eco-ethological In light new arevision systematics moose is out, questions on phylogeny are considered.

Samenvatting

de elandworden artikel De resultaten van onderzoek naar de genetica en de systematiek van in dit gepresenteerd.

onderzoek uit de literatuur Gegevens afkomstig van eigen en over morfologische variatie, erfelijkheid, ecologische de moderne eland worden Veel ondersteunen het idee dat de en gedragskenmerken van besproken. gegevens differentiatiein de moderne Alces een soortspecifiek niveau heeft bereikt, en in twee soorten onderverdeeldkan

de amerikaanse eland worden: de europese elandAlces alces L., voorkomend in Europa, de Oeral en west Siberië, en

A. americanus Clinton, voorkomendin Noord Amerika, oost Siberië en het uiterste oosten van Rusland. Deze soorten verschillenin eland heeft 68 chromosomen in de de amerikaanse eland chromosoomgetal (de europese diploïde set, echter ook in enkele kenmerken. Aan de hand nieuwe wordt revisie 70), maar eco-ethologische van gegevens een ondernomen de worden betreffende hun behandeld. van de systematiek van eland, en vragen fylogenie

Introduction trunk, a strongly developed thorax, a large, elon- and and gated hump-nosed head, a very large

massive and wide upperlip, very large ears, very The moose is the largest representative of the long legs, and a long 'bell', which is a hanging family of . The original English name of Alces outgrowth of skin under the throat. The antlers in alces is 'elk', but nowadays the American name, the male project laterally from the head and are 'moose', is usually used in scientific literature. It usually palmated (shovel-shaped expanded). essentially differs from the other deer by its The front part of the moose torso is more massive habitus: but with rather short very massive a than the hind part (fig. 1).

The of the modern systematic position moose is under discussion. Within their huge Holarctic

habitat, the moose is represented by a number of forms with various degrees of morphological isolation. The opinion prevails that Northern

Eurasia and North America is occupied by one

- Alces alces L. which contains a species -, maximum of eight subspecies: the European

moose or elk (A. a. alces),the East-Siberian moose

(A. a. pfitzenmayeri), the Manchurian or Ussuri

moose (A. a. cameloides), the Buturlin's moose (A.

the a. buturlini), the Alaskan moose (A. a. gigas),

West-Canadian East- moose (A. a. andersoni), the

Canadian moose (A. a. americanus), and the Fig. 1. Moose male. The Kolyma river basin. Photo

Yellowstone a. by Yu. Labutin moose (A. shirasi) (Peterson, 1955, 1974; Sokolov, 1959; Heptner etal., 1961; Wilson &

eland de vallei de Mannelijke in van Kolyma rivier. Reeder, 1992; Geist, 1998). The first description of Foto: Yu. Labutin the moose as a species (Cervus alces) was made by

31 of the modern and review of its The genetics moose a taxonomy

the Karl Linne in 1758, based on the moose from genetic material of the European, Siberian,

and of Sweden as Cervus alces; gradually this name Russian Far Eastern American moose; part well. results has been became used for other moose populations as our already published (Boes- attributed the into the In 1821 Gray species sepa- korov, 1996a, 1996b, 1997, 1998; Boyeskorov,

Since the of the Results from our and rate genus Alces. description 1999). morphological palae- in the American moose as Cervus americanus Clinton ontological studies will appear next two

1822, a number of the researchers considered this issues of Cranium (2004).

from a separate species, distinct the European A. A. alces (Miller, 1899; Seton, 1910; Lydekker, 1915;

1928). An interesting detail is that Anthony, Material and Methods during the20th century several timesmoose were of of from central For the study karyotypes moose imported in England from Canada; some experts and northeast Yakutia (Eastern Siberia), four (Rolfe, 1983) even identified themwith the extinct males and one female were shot in the giant deer, or Irish elk, Megaloceros giganteus. region (Kolyma river basin, , and Unfortunately Linne himself never saw the ; see Boeskorov, 1997). The American moose.

chromosomes were studied in short-term features of the Based some morphological cells of bone with the of upon cultured marrow, use

Flerov (1931, 1934) revealed some diffe- moose, standard methods of staining (Ford & Hamerton,

rences between moose from Europe and North 1956) and silver-staining (Howell & Black, 1980).

and concludedthatEastSiberian moose America, Heterochromatin regions of thechromosomes of latter than are closer to the to European moose. a Kolyma river basin male were revealed by

He moose A. alces proposed two separate species: C-banding (Sumner, 1972). Thepolymorphism of L., Scandinavia, of Eastern inhabiting part DNA markers was studied by Udina (Udina et al, Europe, and Siberia westward of the Yenissey 2002) in seven moose muscle samples collected river, andA. americanus Clinton, inhabiting North by us in Yakutia. America, Siberia eastward of the Yenissey, and with the Far East. Although some scholars agreed this (e.g. Vereshchagin, 1949), Flerov later stated

that the observed differences did not exceed

subspecies level (Flerov, 1952). Nevertheless, his had influence and earlier opinion large many

Russian mammalogists considered the moose

from East and Northeast Siberia to belong to the Gromov American moose (Bobrinsky et al., 1944;

It etai, 1963; Baryshnikov etal., 1981). was agreed

that the American differ upon moose probably

et from European ones at a species level (Gromov

al., 1963; Chernyavsky & Domnich, 1989) or subspecies level (Groves & Grubb, 1982). Subse- quently, differences in chromosome numbers,

and in a number of other genetic, ecological and found between the ethological features were

European and North American moose. For a long

time the morphologic and genetic differences

between the European and American moose

attracted the attentionof the specialists and were

discussed in some monographic works (Filonov, 1983; Bubenik, 1986; Chernyavsky & Domnich, 1989; Danilkin, 1999). Nevertheless, the question

on the number of and of species subspecies male from the south Fig. 2 Karyotype of a moose of modern remains till unsolved. moose now Western Siberia, Altai mountains (after Grafodatsky

& Radjabli, 1988). Inorder to solve this problem we started research

on the systematics, genetics and morphological Het karyotype van een mannelijke eland uit het zui- in 1992. this article variability of the moose In we den van West Siberië, Altai gebergte (naar Grafo- on the differentiation in present our analysis datsky & Radjabi, 1988)

32 CRANIUM 20, 2 - 2003

Siberia and Altai & Genetics of the moose (fig. 2; Grafodatsky Radjabli,

theSaratov district 1985; 1988), (A. Belyanin, pers. Karyological data with 2n=70 also in comm.), and moose occur and Karyological studies of moose in Europe Alaska (Rausch, 1977) (fig. 3; table 1). In addition, the 1960s. These studies northeast North America began in karyotypes of moose from the extreme demonstrated that moose from these regions of Asia (the Kolyma river basin) and from Central differ in chromosomenumber: moose from Scan- Yakutia were found to be identical to those of dinavia and Finland had 68 chromosomes in the American moose (2n=70) (Boeskorov, 1996a, b; diploid set (Aula & Kääriainen, 1964; Nes et al., 1997; Boeskorov et al., 1993) (fig. 4).

1965; Gustavsson & Sundt, 1968), whereas moose The basic number of chromosome arms of all from northeastern Canada and northwestern

= moose is identical (NFa 70). Among the auto- USA had 70 chromosomes (Hsu & Benirschke, somes of European and Altai A. alces two pairs of 1969). These distinctions were until recently bi-armed chromosomes are present, whereas considered A. as an intraspecific dimorphism of North American and East Siberian moose had alces (after Orlov & Bulatova, 1983). Similar only one bi-armed pair. The X-chromosome is results were obtainedfor moose from other parts large and submetacentric in all moose; the of its with 2n=68 geographic range: moose occur Y-chromosome is usually the smallestacrocentric also in Finland (Gripenberg et al., 1986), Western

= the modern of Fig. 3 Geographic differences in moose chromosome numbers (after Boeskorov, 1997. a area

Alces b = the border between the and genus (acc.to: Peterson, 1955, 1974; Filonov, 1983); hypothetical European

= = of American forms of moose; c ”68”, findings of the European chromosomal form; d ”70”, findings the

= East Siberian American chromosomal form; e ”(70)”, chromosome numbers in and North-East moose (data of the author)

Geografische verschillen in chromosoomgetal van eland (naar Boeskorov, 1997). a = huidige verspreidingsgebied

b = tussen de van het geslacht Alces (volgens Peterson, 1955, 1974; Filonov, 1983); de denkbeeldige grens Europe-

= Amerikaanse = de vond- se en vormen van eland; c ”68”, vondsten van Europese chromosomale vorm; d ”70”, sten = in Oost Noordoost van de Amerikaanse chromosomale vorm; e ”(70)”, chromosoomgetallen en Siberische eland (gegevens van de auteur)

33 The genetics of the modern moose and a review of its taxonomy

chromosome. In Finland and Scandinavia a few have expressed pericentromeric C-heterochro- individuals are found with a meta- or submeta- matin blocks that sometimes amount to half and centric Y-chromosome (Aula & Kääriainen, 1964; more of the total chromosomelength. At the same

Gustavsson & Sundt, 1968; Gripenberg et al., time, C-heterochromatin blocks seem to be

1986). lacking on the pairs of bi-armed autosomsand on

the X-chromosome of the Kolyma moose. In the The karyotype of moose from the Kolyma river moose from Finland these blocks are visible on basin has a large amount of C-heterochromatin certain chromosomes (Gripenberg et al., 1986). (Boeskorov et al., 1993) (fig. 4), which is also Moose from Altai hadno heterochromatinblocks characteristic for European and West Siberian on the X-chromosome and the second pair of moose (Grafodatsky & Radjabli, 1985; Gripen- bi-armed autosomes (Grafodatsky & Radjabli, berg et al., 1986). All acrocentric chromosomes Altai has 1985). Thus, it seems that the moose an

intermediate variant of heterochromatin distri-

Table 5 Diploid chromosome sets in moose from bution between European and East Siberian different geographic regions. The number of moose (the latter belongs to the American group). studied is not indicated. The Y-chromosome in all investigated moose is

heterochromatic. Diploide chromosoomparen in elanden van verschil- completely

lende geografische streken. Het aantal bestudeerde The nucleolar organizing regions of chromo- dierenis niet aangegeven. from Yakutia somes (NOR's) in moose are

present in teleomeric regions of the four largest

chromosomes from the and second or third Number of first,

animals Region 2n Reference autosome pairs (fig. 4) (Boeskorov, 1997; Boes-

karyotyped korov etal, 1993). By number and, apparently, on Gustavsson & localization of NOR's the Kolyma and central Sweden 6 68 Sundt, 1968 Yakutian moose do not differ from Finnish A.

+ 68 Nes et al, 1965 Norway alces (Gripenberg et «/., 1986) and Altai moose Aula & Kaari- (Grafodatsky & Radjabli, 1985). A similar number ainen, 1964; and localization of NOR's is found the Finland 218 68 at majority Gripenberg et of the deer of the investigated species genera al, 1986 Rangifer, Cervus and Dama (Grafodatsky et al., , Sara- A.Belyanin 1 68 1990). The stability of this karyotype, which is tov district (pers. comm.) for different of has characteristic genera deer, Western Sibe- Grafodatsky & 5 68 Altai 1985 an and reflect ria, Radjabli, apparently ancient nature, may features of the ancestral of the Eastern Sibe- karyotype family

ria, Boeskorov, Cervidae. 3 70 1997 Central Yaku- data level of differenti- tia Karyological testify a high

be divided chro- North-Eastern ation amongmoose: they can on

= Siberia, mosome number: European (2n 68; Europe, Boeskorov et middle stream 1 70 al, 1993 Western Siberia) and American (2n = 70; North of the Kolyma and river America, Russian Far East Eastern Siberia)

table This of chromosome North-Eastern (fig. 3, 1). dimorphism

lower these for Siberia, numberis not present within two forms, stream of the Our unpublis- 1 70 example, the karyotypes of more than 200 indivi- Kolyma river, hed data duals in Finland all contain 68 chromo- near Chersky exactly somes (Gripenberg et al., 1986).

USA, Alaska + 70 Rausch, 1977 The difference in chromosome number of the Wurster & Be- European and American moose is nirschke, 1967; explained by North-Eastern fusionof 1 70 twoacrocentric autosomepairs into two Canada Hsu & Be- large metacentrics (Grafodatsky et al., 1990; nirschke, 1969 Gripenberg et al, 1986; Groves & Grubb, 1982).

Wurster & Be- Proceeding further from this, it is possible to North-West of nirschke, 1967; assume that European-West Siberianmoose with USA,Montana 1 70 2n=68 from the East Siberi- Hsu & Be- originally originated

nirschke, 1969 an-American moose with 2n=70. Thus, the latter

34 CRANIUM 20, 2 - 2003

should thenbe more ancient, since the karyotype genetic distance between these moose is with 2n=70 is considered ancestral for the family D=0.0602 (Reuterwall & Ryman, 1979; Reuter- Cervidae (Groves & Grubb, 1982; Grafodatsky et wall, 1980), which is less than that between rein- al., 1990). deer(Rangifer tarandus tarandus) andcaribou R. t. caribou ) (D=0.174; Baccus et al., 1983), but larger than that between European red deer (C. elaphus

elaphus) and American wapiti (C. e. canadensis) (D=0.0248; Dratch & Gyllensten, 1985).

Analysis of DNA polymorphism

the last molecular research During years genetic

of moose DNA structure has been carried out.

that have Though in general it was noted moose a low level of genetic variability, interesting results

came out which have a large value for the syste- matics and phylogeny of these animals.

Research of mitochondrial DNA structure of

representatives of all four North American

subspecies of moose has shown the absence of variability of this feature (Cronin, 1992), which

testifies to the of the phenomenon presence "effect of the founder" (Cronin, 1992) in Alces history of North America and confirms the

opinion that the moose moved from its refuge in

Alaska into North America after the last ice sheet

had melted in the beginning of the Holocene (Geist, 1987; Kurten & Anderson, 1980). Fig. 4 Chromosome number of a moose male from

The of a the Extreme North-East of Asia (the Kolyma river study major histocompatibility complex

= of nuclear and basin, near village). a routine staining; b (MHC) through analysis mito-

= C-banding; c = AgNOR-staining chondrial DNA of Scandinavian and Canadian

moose has shown that the low genetic variability Chromosoomgetal van een mannelijke eland uit het is characteristic for them as a whole (Mikko & deel Azië dal meest noordoostelijke van (het van de Andersson, 1995). Theseresearchers investigated rivier, het a= stan- Kolyma vlakbij dorp Zyryanka). the exon-2 of DRB-1 and found variability a gene daard b = = kleuring; C-band patronen; c zilverkleu- distinctions in an allelic set of a locus DRB-1 ring (AgNOR) betweenSwedish and Canadian moose. Among

the revealed 10 alleles of DRB-1, 7 and 4 alleles

Differences in chromosome number between Swedish and Canadian among moose, respecti- factors closely related forms are for subsequent vely, were specific and only one allele was shared genetic isolation (Vorontsov, 1958; Ayala, 1975; between continents. Nevertheless, the variability

Timof et Orlov & Bula- of this be which eeff-Ressovsky al, 1977; gene appeared to low, was

tova, 1983), and they can provide evidence for explained by loss of the variability, probably due differences betweenthese forms. specific to a selection responseon one orseveral preferred

haplotypes MHC, or to a 'bottleneck-effect' Electrophoretic mobility of proteins during evolution, which took place before the The first of blood electrophoretic investigations division into different subspecies. Research of a

in moose showed a absence of proteins complete fragment of a mitochondrial DNA control region determined distinc- electrophoretically genetical (D-loop) has shown distinctions between moose

tions between European and North American from Scandinavia and Canada: mtDNA of the moose et al., 1967; Wilhelmson et (Nadler al., former contained an insertion of 75 nucleotides

1978). (Mikko & Andersson, 1995).

Further electrophoretic research of twenty Recently, the study of DNA polymorphism biochemical loci carried out on a large number of shows the Old a morecomplicated picture among moose from Scandinavia and Canada, revealed a and New World moose. Themost distinctions are low level of differentiation between them. The found between North American and European

35 of The genetics of the modern moose and a review its taxonomy

and the absence of the moose, 75-bp insertion in Ethological features mtDNA in the former is a reliable marker to Sound signalization distinguish them from the latter (studied in Scan- in dinavia, Eastern Europe, Ural and western Theacoustic signals are supposed to be inherited The Siberia). Among moose from Eastern Siberia, the genetically (Nikolsky, 1992). distinctions in the sound between Russian Far East and Manchuria, however, a high signal system and American level of mtDNApolymorphism was found.Many European moose once again that these of specimens (about 45 %) from these regions also evidently testify groups moose are from each other. lack the 75-bp insertion and belong to American essentially diverged haplotype, but theremaining specimens have the Males of European (A. a. alces) and American insertion and belong to the European haplotype moose (A. a. gigas) make different sounds during which differs or to a separate Asian haplotype, the rutting season (Lent, 1974; Bogomolova et al, from the European haplotype in some other 1984). In the European moose, these signals mtDNA features (Hundertmark et al., 2002; resemble grunting, while the Alaskan moose Udina et al., 2002). The absence of Asian-like make deep, resonant sounds followed (or some- haplotypes on the European territory and in times preceded) by nonvocal sound recembling western Siberia corresponds to the geographic suction. Cries of female American moose (A. a. distribution of moose karyotypes (2n=68 and americanus, A. a. gigas), unlike European, are 2n=70). A variable picture similar to the mtDNA longer and probably louder, and have a higher of variants was revealed in thestudy MHC poly- average frequency and demonstrate a wider of moose from Siberia and the morphism Europe, of both and range frequency temporal variations Russian Far East (Udina et al., 2002). As in the (Lent, 1974; Bogomolova et al., 1984). work by Mikko and Andersson (1995), it Among red deer Cervus elaphus two groups of appeared that European and North American subspecies are distinguished on morphological moose differ in their set of DRB1 alleles, but is the with the grounds, as case moose: western, Yakutian and Ussurian three among moose and the or or elaphoid group, eastern, wapitoid alleles typical forA. A. a. americanus and threealleles of group (also known as maral's group). Thelevel typical for Swedish moose were found. These between these divergence groups approaches complicated data show that most DNA diffe- that of the specific divergence, or even reaches it. between and North rences exist European of are also well Representatives these groups American moose. Moose from Eastern Siberia (A. diagnosed on sound signals which males make a. pfitzenmayeri)I and from thesouth of the Far East during the rutting season (Nikolsky, 1984). (A. a. cameloides) represent the highest genetic data variation and probably represent the oldest Helmintological

et moose populations (Hundertmark al., 2002; of helminths At present more than sixty species Udina et this al., 2002), although high genetic known of them (worms) are in moose. Many of the Yakutian and Manchurian moose diversity the occur throughout complete geographic range could well be the result of as ancient hybridiza- of the genus. Others are recorded only in the tion between and American European moose of the of the eastern part geographic range moose (Udina et al., 2002). (Eastern Siberia and the Far East; for example the

cestode Moniezia and the nematodes Similar results on moose mtDNA variation were expansa, Acantospiculum cervipipedis, Parabronema skrjabini, obtained by Lister & Pijlen (1998). The low Spicalopteragia schulzi, and Alcefilaria abramovi). mtDNA variation made them suggest a "star Again others are found in the western of phylogeny" and a recent "bottleneck" effect only part the the fluke moose & 1998; Hundert- Parafasciolopsisgeographic range: among (Lister Pijlen, and the nematodes Setaria labia- mark et al., 2002). Lister & Pijlen (1998) consider fasciolaemorpha, the "to Fasciola aberrant A. a. cameloides be a recently to-papillosa, Bunostomum trigonocephalum,

and reduced rather than relict 1976; Filonov, 1983; derived moose, a hepatica) (Pryadko, Zheleznov, These differences indi- ancestral form", which opinion is not shared with 1990). provide rect evidence for the rather ancient differentiation later studies (Hundertmark et al., 2002; Udina et of the Alces into ai, 2002). genus two large groups.

Pheromones

BetweenNorwegian and North American moose

distinctions in the composition of sexual phero-

mones are found (after Bubenik, 1986).

36 CRANIUM 20, 2 - 2003

Taxonomie diagnostics of A. alces markers (Lister & Pijlen, 1998). The European and bison and A. americanus and their subspe- bison (Bison bonasus) American (B. bison), differ cific systematics being undoubtedly separate species, the DNA at same time only slightly in some Numerous data testify that the differentiation markers (Miyamoto et al., 1989; Morris et al., 1994; modern Alces has reached level a high also among Sipko et al., 1997). The same scenario is re- and that they can be divided into two large vealed between some other closely related European moose (Europe, Ural, Western groups: species of ungulates: C. elaphus and C. nippon, Siberia) and American moose (North America, Capreolus capreolus and C. pygargus, Bos taurusand Eastern Siberia, and the Far East). Representa- B. indicus) (Potapov et al, 1998). Such examples of these differ in chromosome tives groups are not restricted to ungulates. numbers (European moose have 2n=68, In our view, the sum of differences clearly American moose have 2n=70), as well as in demonstrates that the betweenEuro- morphological (pelage colour, body and skull divergence and American moose has reached a specific proportions, non-metrical features of skull and pean level. The clearest evidence for this is the antlers) and some ecological and ethological of chromosome sets. Such features. existence differing a

differentiation between related forms are due to In contrast with these differences, only low levels factors of genetic isolation which, in turn, create of genetic differentiationbetween European and effective post-zygotic factors of reproductive American moose were found in studies of elec- isolation (Vorontsov, 1958; Ayala, 1975; Timo- trophoretic mobility of proteins and DNA feeff-Ressovsky et al, 1977). Modern moose markers (see above). Examples of unequal rates consists thus of two distinct species: Alces alces L. of chromosomal and molecular, morphological of (inhabiting Scandinavia, parts Eastern Europe, evolution however, not rare are, among the Ural and Western Siberia) and A. americanus mammals. There are more cases that show that Clinton (inhabiting North America, Eastern and different ungulate species that can be disting- North-EasternSiberia andpart of theRussian Far uishedby morphological and features yet genetic East). have a low level of genetic distances as deter-

Theforms that are themost far mined by electrophoretic mobility of proteins. genetically spoken removed from each other North low level of diffe- are American For example, a very genetic andScandinavian of Alces. TheSibe- rences was found between the American buffalo populations

rian and Russian Far Eastern moose have (Bison bison) and domestic cattle (Bos taurus), may of between the which to different traces hybridization two species. even belong genera according

This is indicated the Altai A. alces on the basis to the prevailing opinion (Baccus et al., 1983). by of its chromosomeheterochromatin distribution. Among cervids a low level of differentiation Furthermore, the East Siberian and (D=0,149) was foundbetween mule deer (Odocoi- among leus Russian Far Eastern A. americanus some indivi- hemionus) and white-tailed deer (O. virgini- duals are with the European variant of anus) (Baccus et al., 1983), a comparatively low present mitochondrial DNA. genetic distance (D=0,390) was found between

American and sika wapiti (Cervus elaphus spp.) Alces alces Linnaeus, 1758 — European moose, deer & Tate, (C. nippon mantchuricus) (Emerson or elk

1993), etc. On the other side, there are some cases 1758. Cervus alces Linnaeus, Syst. Nat. (10th ed.) 1:16. in which closely related forms (for example, 1827. Cervus coronatus Lesson, Man. .:356. subspecies), which have a low level of D, yet have 1837. Alces machlis Ogilby, Proc. Zool. Soc. London a level of DNA differentiation. For high example, (1836): 135. distance between red the genetic European deer 1843. Alces palmatus Gray, List Mamm. Coll. Brit. (C. elaphus elaphus) and American wapiti (C. e. Mus.: 182. 1860. Alces 4: 86. canadensis) is smaller than that betweenthe Euro- jubata Fitzinger, Naturg. Saugeth.

and American moose Dratch & pean (D=0,0248; Holotype. Unknown. Sweden, Vanersborg. However, these same Cervus Gyllensten, 1985). Characters. Karyotype consists of 68 chromo- elaphus subspecies differ significantly in mito- somes. Colouring of trunk and headrather mono- chondrial DNA markers, therefore it was sugge- tonous, brown and dark-brownish. The lower sted to consider them as two separate species: C. of limbs is whitish. Nasal mirror part light grey, and C. canadensis, based their diffe- elaphus upon (rhinarium) is elongated, ellipsoid. Nasal carti- rences in behaviour and DNA morphology, lages are moderately developed; top of the

37 of The genetics of the modern moose and a review its taxonomy

= Fig. 5 Moose species and subspecies distributionat present. 1 =Alces alees alces, 2 = A. americanus pfizenmayeri, 3

A. 4 = A. 5 =A. 6 = A. ander 7 = A. americanus a. cameloides, a. gigas, a. shirasi, a. soni, a.

Voor de zie Huidige verspreiding van elandsoorten en -ondersoorten. verklaring van nummers, Engelstalig

onderschrift

Till the 20th the contained two muzzle forms an almost straight line, nose height century species is almost equal to width or slightly exceeds it. living subspecies, but at present only one

the remained. Upper (nasal) processes of premaxillary bones are spoon-like widened. Alces alces alces Linnaeus, 1758 - European Description. Large, medium and small-sized elk moose, or whithers' of moose. Mature bulls reach a height

190 200 of 260 275 live 1913. Alces machlis uralensis Matschie, Beroff. Inst. to cm, body length to cm, Jagdkunde. Neudamm2:155."Southern Urals". weight upto 500 or even 600 kg. Comparatively Type specimen in Zoological Museum of short rostrum. Antlers with broad, undivided Humboldt's University, Berlin. single palm or deer-shaped (without palm) 1913.Alces machlis meridionalis Matschie, Beroff. Inst. Antlers about 100 but prevail. spread usually cm, Jagdkunde. Neudamm2:156."Samarskaya reach 135 Width of antler shovel sometimes cm. guberniya". Type specimen in Zoological Museum

is 20 to 30 Antler beam of Humboldt's Berlin. (palmation) cm. compara- University, 1915.Alces alces tymensis Zykowsky, Arch. tively short, usually 5 to 10 cm. Naturg. 9: 42. "Tomsk district, the river". Distribution. of Eastern 80, Tym Type Scandinavia, part specimen in Zoological Museum of Humboldt's Europe, Ural, Western Siberia (including Altai) University, Berlin (?). eastward approximately till the river. Yenissey 1915.Alces alces alces Lydekker, Cat. Ungulate Mamm. Brit. Mus. (Nat. Hist.) 4: 232.

38 CRANIUM 20, 2 - 2003

Holotvpe. Unknown. Sweden, Vanersborg. somes. Colouring of trunk and head mainly

Characters. Upper processes of the premaxillary brown and dark-brown. Different parts of the

bones are widened spoon-like, and do not come body differ in colour from light-brown toalmost

into contact with the nasal bones. black. The colour of the lower part of the limbs medium-sized from Description. A large to moose. ranges light-brown to dark-brown or grey.

Antlers with broad undivided deer-like Nasal widened in the and palm or mirror is upper part

antlers prevail. Condilobasalskull length usually pear-shaped. Nasal cartilages are strongly deve- of the muzzle 530 to560 mm. loped; top is humped, nose height

Remarks. In spite of the fact that West Siberian exceeds sufficiently its width (almost 1,5 times). of the bones moose are a little larger than East European Upper processes premaxillary are do differ narrowed. moose, they not significantly. wedge-like Distribution. Scandinavia, Finland, Poland Description. Large, medium and small-sized Czechia and bulls reach withers' of (whence it comes into East Germany, moose. Mature a height Baltic Russia of 220 and of 300 and Austria), countries, (wood zones cm more, body length cm

the European part, Ural and Western Siberia more, live weight to600 kg and more. Comparati-

(including Altai), north of Kazakhstan) and the vely long rostrum. Antlers with broad, divided western part of Mongolia (fig. 5). (double) palm prevail; anterior (brow) palm is separated from the mainpalm. Only for Ussurian Alces alces caucasicus Verestschagin, 1955 - moose the deer-like antlers are characteristic. Caucasian moose Antlers 100 sometimes spread usually more cm, 1955.Alces alces caucasicus Zool. with recorded of Verestschagin, even 150to 160 cm, a maximum Journal 34, 2: 461. 200 cm. Width of antler shovel (palmation) is 20 to 1956. Alces alces caucasicus Scientific Naniev, papers 40 cm. Antler beamcomparatively long, usually 9 of the North Osetian State Pedagogical Inst. 20. to 11 cm.

ZIN 23615 skull withoutlower Lektotype. (male), Distribution. Forest zone of North America, Northern Osetia, Makhchet district, jaw. Digo- Eastern Siberia west-ward to the Yenissey river rized Urukh river. ZIN cave, RAS, St.-Petersburg. basin, Russian Far East, north andeast Mongolia, Remarks. extinct the of the 20th Got at beginning north-eastern China. century, so information on this moose is based on Remarks. Among American moose upto seven the bone material only. subspecies were described: three in Eurasia and Characters. of the Upper premaxillary four North with six processes in America. We agree subspe- bones are wide and and spoon-like, very long cies, and consider moose from the extreme

reach the nasal bones. Antlers are practically North-East of Asia and Alaska consubspecific deer-like. with A. a. gigas. The treatmentof four subspecies

Medium and small-sized moose. Description. in North America is rather traditional, and it is

Condilobasal skull is 520-540 mm. length more likely that there are not sufficient diffe- Distribution. Forests of the foothills of the Nort- rences betweenA. a. americanus and.A. a. andersoni hern Caucasus, of Transcaucasia of part (shore to treat them as two subspecies (Grubb & the Black Sea). Gardner, 1988). Nevertheless we have not enough cranial material of these subspecies, and Alces americanus Clinton, 1822 — American

therefore consider them for the time as moose being

separate subspecies. 1822. Cervus americanus Clinton, Letters on Nat. Hist, and Inst. Resources of : 193. A. americanus includes the following six subspe- 1835.Alces americanus Jardine, Nat. 21 Library cies, which are considered according to their (Mammalia: deer, antelope, camels, etc.): 125. geographical distribution in Eurasia and North 1867. Cervus cameloides Milne-Edwards, Ann. Sci. America (fig. 5): Nat., Zool. 7: 377.

1899. Alces Miller, Biol. Wash. 13: 57. gigas Proc. Soc. Alces americanus pfitzenmayeri Zykowsky, 1902. Alces bedfordiae Lydekker, Proc. Zool. Soc. 1910 — East Siberian, or Yakutian moose London 1:109.

Alces 1910. Alces pfizenmayeri Zykowsky, Wild und Hund 1910. pfitzenmayeri Zykowsky, Wild und Hund

16: 807. 16: 807.

1911. Alces machlis yakutensis Millais, The Field 118: Unknown. north of White- Holotype. "Country 113. stone" (probably in western Adirondak region), Holotype. Unknown. Eastern Siberia, Yakutia, New York. the Aldan river basin. Characters. Karyotype consists of 70 chromo-

39 The genetics of the modern moose and a review of its taxonomy

Characters. Antler palmation is divided into two subspecies in Eurasia. Antlers are deer-like

parts, the anterior (brow) palm is divided into shape, without palmation. Antler length is 80 to

one to three branches. Antler length is 90 to 110 90 cm. Condilobasal skull length is 505 to545mm.

Condilobasal skull is 540 Small- medium-sized In cm. length usually to Description. to moose.

565 mm. Primorie region mature bulls reach a withers'

A bulls of 170-190 of 230 280 Description. large-sized moose. Mature height cm, a body length to

a withers' of 190 to 205 live of 250 to 400 with an of reach height cm, body cm, weight kg average of length of 265 to 290 cm, live weight 400 to 500 300 kg. In Manchuria this moose may be smaller.

Antlers to 110 width of antler The of two males from Great kg. spread 100 cm, average body length is 20 30 antler is 230 withers' of shovel (palmation) to cm, length Qingan was cm at a height 165,5

usually approximately 100 cm. cm (Wang & Liu, 1989). A dark colour for body Distribution. Eastern Siberia west-ward to the and head is characteristic. Antlers spread 80 to

river northeast-ward the 100 adult males have three five tines. Enissey basin, to cm; to Chersky ridge, Western Transbaikalia, Northern Rostrum is comparatively short. Mongolia, partly Manchuria (?) (fig. 5). Distribution. The south of the Russian Far East (Shikhote Alin), North-Eastern China Alces americanus cameloides Milne-Edwards, (Manchuria) and the extreme east of Mongolia 1867 — Ussurian (Manchurian) moose (Great Qingan) (fig. 5). In the adjacent territories

1867. Cervus cameloides Milne-Edwards, Ann. Sci. (Lower Amur river, Eastern Transbaikalia) and Nat. Zool. 7: 377. the sometimes in main area hybrids with A. a. 1902. Alces bedfordiae Lydekker, Proc. Zool. Soc. pfizenmayeri were noted. They usually have large London 1:109. body sizes and deer-like antlers. 1928. Alces alces bedfordi "Lydekker" Bobrinskoi, Field Guide to Game Animals: 95. Ussuriysky Alces americanus gigas Miller, 1899 — Alaskan (Primorsky) kray (region). Specification of the name moose and type locality for Alces bedfordiae Lydekker. 1940. Alces alces cameloides China and Allen, Mamm. 1899. Alces gigas Miller, Proc. Biol. Soc. Wash. 13: 57. Mongolia, Am. Mus. Nat. Hist. 11, 2:1205. 1904.Alces americanus gigas Osgood, N. Am. Fauna,

24: 29. Holotype. Unknown. China, Heilonjiang 1915. Alces alces gigas Lydekker, Cat. Ungulate ("Manchuria"). Mamm. Brit. Mus. (Nat.Hist.) 4: 237. Remarks. Till for moose from the south of present 1982. Alces alces buturlini Chernyavski et Zhelesnov, the Russian Far East North-Eas- (Shikhote Alin), Bulletinof the Moscow Society of Naturalists. Biol. tern China and the extreme east of Mongolia, Department 87, 5: 28. which all belong to one and the same subspecies, Holotype. No 86166, male. "North side of Tustu- subspecific names as A. cameloides, andA. bedfordi mena Lake, Kenai Peninsula, Alaska" (U.S. used. Alces described in 1902 are bedfordiae was by National Museum).

based on a skull with deer-like antlers. Lydekker Remarks. A time it long ago was already This material probably originated from Eastern supposed that the largest moose with antlers Siberia. In 1928 Bobrinsky partly changed the characterised by a huge shovel, living on both of of name this form moose andhas specified the sides of the Bering strait (in the extreme Nort- type locality: Ussuriysky kray (Ussurian region), heast of Asia and in Alaska) belong to one and the but this change and specification have no value same subspecies: A. a. gigas (Kulagin, 1932; (Heptner et al., 1961), andA. a. bedfordiae andA. a. Yegorov, 1972). However, others (Chernyavsky i can be more than for bedfordi nothing synonyms & the Eurasian Zheleznov, 1982) assigned popu-

A. a. cameloides. lation to a separate subspecies (A. a. buturlini), Some features of this morphological moose are based their and skull upon larger body size in closer to thoseof moose European (short rostrum, Alaskan A. which respect to a. gigas, really are deer-like antlers) but other diagnostic features smaller in many parameters (Peterson 1955; (narrow of of edges upperprocesses premaxillary Franzmann et ai, 1978). From the middle of the dark bones, body pelage, grey or light-brown 20th century onwards a decrease took place in the colour of legs) show that Manchurian moose size of A. a. gigas because of intensive hunting belong to the American group. The karyotype of pressure (Chernyavsky & Domnich, 1989). this is not known but most subspecies yet, Earlier sizes of not very large Alaskan moose, it belongs to the American species probably smaller than A. a. buturlini, were noted: withers' based markers of upon some molecular-genetic and height to 230 cm, a living weight of 800 kg DNA & Danilkin, 1998; Udina etal., 2002). (Udina (Seton, 1910; Stone, 1924 - after Peterson, 1955;

Characters. Smallest and most dark-colored Anthony, 1928). Chernyavsky & Zheleznov

40 CRANIUM 20, 2 - 2003

thanA. (1982), comparing data of a small sample of skulls thanA. A. a. americanus and narrower a. gigas

of the Yukon river and from the upper stream (data A. a. shirasi) (after Peterson, 1955).

given by Youngman, 1975), noted that their sizes Description. A large to medium-sized moose. of from bulls reach withers' of 180 200 were smaller than those moose Kolyma Mature a height to

of 255 275 live of andAnadyr region. Our craniometric parameters cm, body length to cm, weight

of from the Northeastof Asia and 350 550 Antlers 90 115 moose extreme to kg. spread usually to cm,

show that have similar width of antler shovel 20 to 30 Alaska they very parame- (palmation) cm,

ters (Boeskorov, 1998; Boeskorov & Puzachenko, number of tines 8 to 12. Condilobasal skull length

2001). This allow us to consider the subspecies 545 to 580 mm.

A. a. buturlini invalid. In our view they belong Distribution. Northern Michigan and Minnesota,

with the moose of Alaska and the Yukon river western Ontario, westward to central British

basin to the subspecies A. a. gigas. Columbia, north to eastern Yukon Territory and Characters. Very largeantlers with long andwide Mackenzie Delta, Northwest Territories. In the of with palmation separated into two parts. Anterior northwestern part its area hybrids A. a. divided four branches. which thereforehave (brow) palm is in to six gigas occur, larger sizes (fig.

number of tines antler: ten to fifteen. Large per 5).

Antler length usually exceeds 110 cm. Condilo- Alces americanus shirasi Nelson, 1914 — Yel- basal skull length is 570 to 600 mm. lowstone moose Description. Largest living moose. Mature bulls of 1914. Alces americanus shirasi Nelson, Proc. Bio. Soc. usually reach a withers' height 200 to220 cm or Wash. 27: 72. 280 330 live more, body length is to cm, weight 1952. Alces alces shirasi Peterson, Roy. Ont. Mus. 500 to 600 kg or more. Representatives of the Zool. and Paleo., Contrib. 34: 23. modern Alaska-Yukon population are smaller in No 202975. "Snake Lincoln size (size decrease took place due to intensive Holotype. River, U.S. National Museum. hunting) than those of the Eurasian population. County, Wyoming". Characters. the smallest of all North Nevertheless, the former are the largest subspe- Probably

American moose. Nelson noted: "A cies inNorth America. Mature bulls usually reach (1914) medium form with nasal aperture relatively a withers' height of 190to 210 cm, body length is of 270 320 live 400 600 Antlers wide. Colouration pelage the back to cm, weight to kg. along than other North 120 150 with maximum 200 averages paler American spread to cm, a upto width of antler shovel 30 40 forms". However, Bubenik's data (1986) do not cm; (palmation) is to confirm the coloration, and the cm. paler systematic

status need revision. Distribution. In North America: wood areas of may

A medium-sized moose. Mature Alaska, western Yukon, northwest of British Description. bulls reach of 255 275 and Columbia. Extreme Northeast of Asia: basins of a body length to cm, a live of 300 450 have the rivers Indigirka, Kolyma, Anadyr, Penzhina. weight to kg. They relatively small hoofs. Antlers are probably the smallest in In the recent past (down to the beginning of the North America. Condilobasal skull is 535 20th century) they were distributed more widely: length

the the south to 550 mm. in west up to river, in the upto Distribution. Western and the Aldan river (fig. 5). Wyoming, eastern northern Idaho, western Montana, northward Alces americanus andersoni Peterson, 1950 — into southwestern and southeastern

Northwestern moose, or West Canadian moose British Columbia (fig. 5).

1907. Alces columbaeLydekker, Field 9:18. Alces americanus americanus Clinton, 1822 — 1950. Alces americana andersoniPeterson, Roy. Ont. Eastern or East Canadian moose Mus. Zool., Occ. Pap. 9:1. moose, 1952. Alces alces andersoniPeterson, Ont. Mus. Roy. 1822. Cervus americanus Clinton, Letters on Nat. Hist, Zool. and Paleo., Contrib. 34: 24. and Inst. Resources of New York: 193.

1901. Alces machlis americanus Lydekker, Great and Holotype. No 20068, male, complete skeleton and Small Gameof Europe, etc.: 46. skin. "Section 27, Township 10, Range 16, Spruce- 1924. Alces americana americana Miller, U.S. Natl. wood forest reserve mi. E. Mani- (15 Brandon), Mus„ Bull. 128: 490. toba". Ontario Museum of and Royal Zoology 1952. Alces alces americana Peterson, Roy. Ont. Mus. Palaeontology. Zool. and Paleo., Contrib. 34: 28.

Characters. Differ from other North American

forms chiefly in cranial details, especially with Holotype. Unknown. "Country north of White- respect to the shape of the palate (relatively wider stone" (probably inwestern Adirondack region),

41 The genetics of the modern moose and a review of its taxonomy

differentiation of the American moose NewYork. stronger and Characters. Narrow palate relatively to length of (five or six subspecies in Eurasia North than of the also testi- tooth row Peterson, moose upper (after 1955). America) European Description. A large to medium-sized relatively fies the more ancient origin of the former.

darkmoose. Maturebulls reach withers' height of Based the lack of variation in mitochondrial upon 180 190 of 250 290 live to cm, body length to cm, DNA restriction fragments in North American weightof 350 to 450kg. Antler spread is about 100 moose it can be concluded that this population width of antler 20 30 cm, palmation to cm, mustbe and that entered relatively young moose number of tines seven to nine. Condilobasal skull North America from Northeast Asia during the length is 540 to 560 mm. late Wisconsinan (Cronin, 1992). At the same time Remarks. Probably there are no significant diffe- from Little there is a fossil yerling bull mummy rences between A. a. americanus and A. a. ander- Eldorado Creek, Alaska (AMNH, AM 274-4002), soni; they may represent one subspecies. which radiocarbon dated 33.000 BP is y. (after Distribution. From and west- This Guthrie, 1990b). mummy probably belongs ward through to central Ontario (fig. 5), Cervalces to Cervalces latifrons or to stag-moose sp. where it apparently intergrades with A. a. ander- (Guthrie, 1990a, 1990b), but more likely to true soni. Introduced into Newfounland, where it is Alces This makes it moose, (Boeskorov, 2001). now established (after Bubenik, 1986). that inhabited Alaska possible true moose more

than thousand nevertheless, we thirty years ago, consider that the time whenAlces penetrated into Conclusion North unclear. America is till now In our opinion

the differentiation of modern North-American Analysis of karyological data indicates that the

moose into three or four could not forms from which later the European-West-Sibe- subspecies have taken place during ten thousand For rian and theEast-Siberian-American moose origi- years. example, the two amphi-Beriangian populations nated, probably stood apart already at the Late

of A. a. still have not reached a Pleistocene. The level of differentiation between gigas subspecific

level of differentiation their ten modern and American based notwithstanding European moose, thousand of isolation from markers of has years morphological upon molecular-genetic DNA, each other (Boeskorov, 2001). allowedtoestimate the timeof their divergence at

than 100 thousand & more years ago (Mikko

Anderson, 1995) or somewhere between 75 and

et 150 thousand years ago (Udina ai, 2002). Thus, Acknowledgments the divergence of Alces into two species appa- for The author thanks the following persons rently occured at the end of the Middle - begin- discussion of the material and valuable remarks: ning of the Late Pleistocene. Which species of Prof. N.N. Vorontsov, Dr. N. A. Formozov, Dr. I. is the difficult the moose mostancient is to say on A. Vislobokova, Dr. E. A. Lyapunova, Prof. A. A. basis of paleontological finds. The presumably Nikolsky, Dr. P .A. Nikolsky, Dr. A. V. Sher, and earliest remains of late Middle Pleistocene true Dr. I. Yu. Baklushinskaya. The author is deeply found from to moose are Western Europe up grateful for help with the collections to Dr. J. Eastern Siberia. At the time data same genetic Alexander, Mr. A. K. Antonov, Dr. G. F.

testify that the East-Siberian - American moose Baryshnikov, Dr. Yu. M. Dunishenko, Prof. K. most likely are more ancient than European Heissig, Dr. R. Hiitterer, Dr. U. Joger, Dr. R. Kraft, The with 2n=70 is considered moose. karyotype Dr. V. F. Lyamkin, Dr. R. MacPhee, Mr. J. Opper- as ancestral for the family Cervidae; the similar mann, Dr. I. Ya. Pavlinov, Prof. G. Storch, Prof. M. chromosomal set of A. americanus is therefore Stubbe, Dr. A. N. Tikhonov, andDr. E. I. Zholner- the archaic which more probable set, was trans- ovskaya. We are extremely grateful to Dr. Yu. V. formed into the 2n=68 chromosomal karyotype Labutin, Dr. Ya. L. Volpert, Mr. 1.1. Semenov, and of A. alces as a result of Robertsonian fusions others who helped us during the collecting of (Groves & Grubb, 1982; Grafodatsky et ai, 1990). material in the field. Finally, the author brings In Eastern Siberia and the Russian Far East the cordial gratitude to his wife M. V. Shchelchkova largest variety of haplotypes of mitochondrial for her constant work and the help in my editing DNA is revealed, which might indicate that this of the manuscript. region was the closest to the place of origin and evolution of modern Alces (Hundertmark et al.,

2002; Udina et al, 2002). In our opinion, the

42 CRANIUM 20, 2 - 2003

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