CRANIUM 20, 2 - 2003 of the modern and review of its The genetics moose 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 deer. 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 Sakha giant deer, or Irish elk, Megaloceros giganteus. region (Kolyma river basin, Gorny district, and Unfortunately Linne himself never saw the Khangalassky district; 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