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3 2 Kapitanova.Pm6 Russian J. Theriol. 3 (2): 89106 © RUSSIAN JOURNAL OF THERIOLOGY, 2004 Cranial morphometry and taxonomy of argali, Ovis ammon (Artiodactyla, Bovidae), from former Soviet Union and Mongolia Darya V. Kapitanova, Alexey V. Lopatin, Andrey E. Subbotin & William A. Wall ABSTRACT. A partial revision of the subspecies taxonomy of argali Ovis ammon was done on the basis of cranial features using the modern statistical methods (discriminant analysis, cluster analysis, multidimen- sional scaling). Skulls of mature males from different populations of the former Soviet Union and Mongolia were studied. A system of measurements and characters was developed for the thorough description of the skull and its bones. Three discrete cranial types were distinguished: Nuratau (O. a. severtzovi), Kyrgyz- Kazakhstan (O. a. nigrimontana, O. a. polii, and O. a. karelini) and Mongolian (O. a. ammon and O. a. darwini). A clear isolation of O. a. severtzovi from other argalis was noted, as well as from moufflons and urials. Differences between O. a. collium and O. a. karelini do not attain the subspecies level; therefore, these forms are combined into the subspecies O. a. karelini. O. a. ammon and O. a. darwini form a group isolated from other argalis and differentiated into two subspecies based on the discriminant analysis. KEY WORDS: Ovis ammon, cranial morphometry, subspecies level taxonomy, Kyrgyzstan, Kazakhstan, Tajikistan, Uzbekistan, Mongolia. Darya V. Kapitanova [[email protected]], Severtzovs Institute of Ecology and Evolution, Russian Academy of Sciences, Leninsky pr. 33, Moscow 119071, Russia; Alexey V. Lopatin [[email protected]], Paleontological Institute, Russian Academy of Sciences, ul. Profsoyuznaya 123, Moscow 117647, Russia; Andrey E. Subbotin [[email protected]], Severtzovs Institute of Ecology and Evolution, Russian Academy of Sciences, Leninsky pr. 33, Moscow 119071, Russia; William A. Wall [[email protected]; [email protected]], Safari Club International Foundation, 501 2nd Street, NE, Washington, D.C. 20002, USA. Êðàíèàëüíàÿ ìîðôîìåòðèÿ è òàêñîíîìèÿ àðõàðîâ Ovis ammon (Artiodactyla, Bovidae) òåððèòîðèè áûâøåãî ÑÑÑÐ è Ìîíãîëèè Ä.Â. Êàïèòàíîâà, À.Â. Ëîïàòèí, À.Å. Ñóááîòèí, Ó.À. Óîëë ÐÅÇÞÌÅ. Ïðîâåäåíà ÷àñòè÷íàÿ ðåâèçèÿ ïîäâèäîâîé ñèñòåìàòèêè àðõàðà Ovis ammon íà îñíîâå êðàíèîëîãè÷åñêèõ ïðèçíàêîâ ñ èñïîëüçîâàíèåì ñîâðåìåííûõ ìåòîäîâ ñòàòèñòèêè (äèñêðèìèíàíò- íûé àíàëèç, êëàñòåðíûé àíàëèç, ìíîãîìåðíîå øêàëèðîâàíèå). Èññëåäîâàíû ÷åðåïà ïîëîâîçðåëûõ ñàìöîâ èç ðàçíûõ ïîïóëÿöèé ñ òåððèòîðèè áûâøåãî ÑÑÑÐ è Ìîíãîëèè. Ðàçðàáîòàíà ñèñòåìà ïðîìåðîâ è èíäåêñîâ, äåòàëüíî îïèñûâàþùèõ ñòðîåíèå ÷åðåïà è ñîñòàâëÿþùèõ åãî êîñòåé. Âûäåëå- íû òðè äèñêðåòíûõ êðàíèàëüíûõ òèïà: íóðàòèíñêèé (O. a. severtzovi), êèðãèçñêî-êàçàõñòàíñêèé (O. a. nigrimontana, O. a. polii è O. a. karelini) è ìîíãîëüñêèé (O. a. ammon è O. a. darwini). Îòìå÷åíà ÷åòêàÿ îáîñîáëåííîñòü áàðàíà Ñåâåðöîâà êàê îò äðóãèõ ïðåäñòàâèòåëåé ãðóïïû àðõàðîâ, òàê è îò ìóôëîíîâ è óðèàëîâ. Ðàçëè÷èÿ ìåæäó O. a. collium è O. a. karelini íå äîñòèãàþò ïîäâèäîâîãî óðîâíÿ, ïîýòîìó ýòè ôîðìû îáúåäèíåíû â îäèí ïîäâèä O. a. karelini. O. a. ammon è O. a. darwini îáðàçóþò õîðîøî îáîñîáëåííóþ îò äðóãèõ àðõàðîâ ãðóïïó, ïî ðåçóëüòàòàì äèñêðèìèíàíòíîãî àíàëèçà äèôôåðåíöè- ðîâàííóþ íà äâà ïîäâèäà. ÊËÞ×ÅÂÛÅ ÑËÎÂÀ: Ovis ammon, êðàíèàëüíàÿ ìîðôîìåòðèÿ, ïîäâèäîâàÿ ñèñòåìàòèêà, Êèðãèçñòàí, Êàçàõñòàí, Òàäæèêèñòàí, Óçáåêèñòàí, Ìîíãîëèÿ. Introduction size. The geographical confinement of habitats was the main criterion for the separation into species and subspe- The taxonomy of wild sheep (genus Ovis L.) has cies. The results of these studies are contradictory and been developed since the late 19th century. The works of reflect the development from the typological to polytypic Severtzov (1873), Lydekker (1898), Nasonov (1923), concept of mammal species (Vorontsov et al., 1972). No and Tsalkin (1951) are of fundamental value. The sys- clear idea has been proposed up to now. The position of tems of these scientists were mainly based on the differ- one of the above researchers is usually followed in the ences in cranial proportions, the direction of horn twist- present-day works on the mammal taxonomy. ing, and some other external morphological features The study of chromosome sets in the geographical such as wool color, presence of a ruff, and total body forms of sheep revealed four genome types in the group: 90 D.V. Kapitanova, A.V. Lopatin, A.E. Subbotin & W.A. Wall 52-chromosome snow sheep (O. nivicola), 54-chromo- the species O. ammon, among moufflons (Tsalkin, 1951; some moufflons (O. musimon and O. orientalis) and Sokolov, 1959; Heptner et al., 1961; Vorontsov et al., North-American bighorn sheep (O. canadensis and O. 1972). After a thorough study of its horns, Ishunin dalli), 56-chromosome argalis and arkhars (O. ammon), (1968) supposed that the Severtzov's argali is closer to and 58-chromosome urials (O. vignei) (Vorontsov et the Karatau sheep than to the western strains. Valdez al., 1972; Korobitsyna et al., 1974). However, some (1982) ranked the Severtzov's argali among urials. Sopin species differing in chromosome sets can be ranked (1982) did not consider the severtzovi form in the among subspecies, because data are available that the analysis of argali subspecies, apparently placing it among reproductive isolation of 54-, 56-, and 58-chromosome urials. More recently, Sopin (1984) concluded that the wild sheep is evidently imperfect, if any (Orlov, 1978). In Severtzov's argali is much closer to Karatau sheep this approach, the genus Ovis includes two species (O. (nigrimontana form) than to urials and ranked it among ammon and O. canadensis), which in turn include sub- argalis as a subspecies. The study of the Severtzov's species with different numbers of chromosomes (Orlov, wild sheep karyotype undoubtedly showed that it be- 1978). longs to the so called arkhar-argaloid group of sheep, These disagreements called for the revision of the Ovis ammon s. str., although, in macromorphological subspecies taxonomy of the group and the establish- terms, the Severtzov's sheep occupies an intermediate ment of taxonomic ranks of geographical forms better place between urials and argalis (Lyapunova et al., 1997). defined in morphological terms. Sopin (1982) analyzed When studying the geographical diversity of argalis 13 parameters describing the cranial proportions and in the Mongolian and Russian Altai regions, Sopin & some bones for a series of adult argali males. Along with Sukhbat (1980) noted significant differences in the skull craniological features, color variation was also consid- size and its variation between these two populations. ered. Sopin confirmed the subspecies rank of O. a. Nonetheless, they concluded that the populations are nigrimontana, O. a. ammon, O. a. darwini, O. a. ko- morphologically close and that the noted differences are zlovi, O. a. dalailamae, and O. a. hodgsoni and com- of population rank. Later on, these authors studied the bined three forms O. a. karelini (Tien Shan), O. a. polii populations of Mongolian and Gobi Altai for determin- (Pamirs) and O. a. collium (Kazakhstan hillocky area) ing their taxonomic ranks (Sopin & Sukhbat, 1983). into the subspecies O. a. polii. The Mayr coefficients of They revealed reliable differences in skull sizes (Mon- discrimination (calculated as the ratio of the difference golian Altai argalis are bigger than the sheep of Gobi between the mean values of a parameter in the samples Altai) and cranial proportions (Gobi Altai argalis have compared to the sum of standard deviations) were used shorter and narrower skulls). Therefore, a conclusion as an analytical tool. However, Mayr (1969) noted that about the subspecies status of both populations was coefficients of discrimination give only approximate drawn. estimates, because their calculation is based on inexact Valdez (1982) postulated the existence of six argali assumptions (e.g., that the distributions are normal and subspecies: O. a. ammon (=O. a. mongolica, O. a. that statistic characteristics of a sample are equal to jubata), O. a. nigrimontana, O. a. darwini, O. a. hodg- those of the whole population). This calls into question soni (=O. a. blythi, O. a. brookei, O. a. dalailamae, O. the results obtained. Most authors are inclined to con- a. adametzi), O. a. polii, and O. a. karelini (=O. a. sider O. a. polii as an argali subspecies well distin- heinsii, O. a. collium, O. a. sairensis, O. a. littledalei, guished from O. a. collium and O. a. karelini, in spite of O. a. humei). the hybridization zone existing between O. a. polii and Geist (1991) analyzed differences between the geo- O. a. karelini at the boundary of their ranges (Nasonov, graphical forms of argalis from the nuptial coat of males 1923; Rumyantsev, 1935; Tsalkin, 1951). As for the (wool color during the rutting season). In a thorough karelini and collium forms, many authors consider them review of works on the taxonomy of this group, Geist very close to each other (Tsalkin, 1951; Heptner et al., made criticisms on the procedure used; e.g., he indicates 1961; Sludskii et al., 1983): the only differences in the the absence of statistical analysis of the results, incom- premaxilla length, which is longer in the collium form plete description of the geographical forms, and inexact (Tsalkin, 1951), and in the character of horn twisting gridding of specimens. Geist divides the species into (Nasonov, 1923) were noted. seven subspecies: O. a. nigrimontana, O. a. hodgsoni, Of great interest is the taxonomic position of Sever- O. a. jubata, O. a. darwini, O. a. ammon, O. a. polii and tzov's argali inhabiting the Nuratau and Aktau ranges, O. a. karelini. the southeastern Kyzyl-Kum Desert. Almost all authors It is noteworthy that Valdez and Geist combine the note the intermediate morphological position of this collium and karelini forms. form between argalis and urials. Nasonov (1923) de- Fedosenko (2000) states the existence of eight sub- scribed this form as an O. polii subspecies, although he species: O. a. polii (Pamir or Marco Polo argali), O. a. noted its traits in common with sheep of the group collium (Karaganda or Kazakhstan argali), O. a. kareli- Mufloniformes. In later investigations, the severtzovi ni (Tien Shan argali), O.
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