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Ellobius cf. barbarus Pomel, 1892 MATÉRIEL. — N = 27+; deux M1 G, une M1 D, deux M3 G, une M3 D, cinq m1 G, trois m1 D, deux m2 G, deux m2 D, six m3 G et trois m3 D isolées (+ fragments d’émail de molaires isolées non identifiables). DESCRIPTION ET COMPARAISONS Ellobius Fischer, 1814 est le seul Arvicolinae ayant jamais existé au Maghreb, et seulement au cours du Pléistocène moyen (Stoetzel 2013). Venu du Proche-Orient, son histoire évolutive en Afrique du Nord diffère de celle des populations asiatiques. Se succèdent ainsi plusieurs espèces au cours du Pléistocène moyen: E. africanus Jaeger, 1988, E. atlanticus Jaeger, 1988, E. barbarus Pomel, 1892, E. zimae Jaeger, 1988, avant de disparaître au Pléistocène supérieur. Certains auteurs suggèrent que le genre Ellobius aurait pu s’éteindre plus tôt au Maghreb occidental (Maroc et ouest de l’Algérie) qu’au Maghreb oriental (Tunisie) (Mein & Pickford 1992). De plus, sa présence à Tighennif, dont l’âge serait potentiellement à reculer considérablement d’après des études récentes (Sahnouni et al. 2018b; Pickford 2020), pourrait également refléter une présence au Maghreb dès le Pléistocène inférieur (à confirmer avec l’analyse du matériel microfaunique des fouilles récentes; Saidani, thèse en cours). Trois de ces espèces représenteraient une lignée anagénétique: E. africanus – E. atlanticus – E. barbarus. Au cours du temps, on observe une augmentation progressive de la taille des molaires (longueur et largeur), une augmentation de l’indice d’hypsodontie, un développement d’échancrures d’émail, l’angle rentrant lingual 1 des M2 devient plus profond et plus large et le lobe antérieur des m1 se simplifie (Jaeger 1975, 1988). E. zimae serait issu d’une autre lignée (s’enracinant peut-être chez E. africanus) et présente des caractères dentaires un peu distincts, comme la disparition de la racine linguale sur les M1, des m3 réduites, des m1 à deux racines avec un lobe antérieur large et court et un moindre développement des échancrures d’émail (Jaeger 1988). Au sein de la lignée E. africanus – E. atlanticus – E. barbarus, E. africanus est le plus petit, l’indice d’hypsodontie est faible, les molaires inférieures sont plus étroites, le lobe antérieur de la m1 est large et complexe avec de nombreux replis d’émails, les échancrures d’émail sont peu développées, les M2 et m2 sont nettement moins longues que les M1 et m1. E. atlanticus a une taille et un degré d’hypsodontie intermédiaires entre E. africanus et E. barbarus, le lobe antérieur de la m1 est court mais large (étendu du côté lingual), lisse et dépourvu de plissements d’émail, les échancrures d’émail sont peu développées, les M2 sont presque aussi longues que les M1. E. barbarus est le plus grand (molaires plus longues et plus larges), l’indice d’hypsodontie est plus important que chez les autres espèces fossiles (mais inférieur aux espèces actuelles), les échancrures d’émail sont relativement élevées, le lobe antérieur de la m1 est simple, étroit et allongé mais porte parfois un ou deux plissements d’émail sur le rebord antéro-labial, la racine linguale de la M1 est réduite, les M3 sont plus réduites (Jaeger 1988; Geraads 1994, 2016a). Chez les représentant asiatiques actuels (E. lutescens Thomas, 1897, E. fuscocapillus Blyth, 1843, E. talpinus Pallas, 1770), la taille est plus réduite, le degré d’hypsodontie est plus important, les échancrures d’émail sont plus élevées, la proportion des M2 et m2 est moindre, sur la m1 le lobe antérieur est large et plus simple (pas d’incisions labiales ou linguales) et des différences existent également au niveau de l’ouverture des angles rentrants linguaux et de la position respective des triangles (Jaeger 1988). Au moins deux m1 de Ben Kérat (Fig. 11) présentent plusieurs indentations sur leur lobe antérieur (la plupart des dents sont abimées sur leur partie antérieure), un caractère qui est habituellement plutôt associé aux formes anciennes (comme E. africanus), bien qu’il se retrouve aussi parfois chez E. barbarus (Jaeger 1988). De plus, l’indice d’hypsodontie observé dans le matériel de Ben Kérat est assez modéré (moy 0,83; min0,73; max 1,02), se rapprochant davantage de E. africanus (0,92 à Tighennif) ou E. atlanticus (1,03 à la carrière Thomas) que de E. barbarus (1,14 à Aïn Mefta) (Jaeger 1988; Table 7). Il faut cependant garder à l’esprit que la valeur de l’indice d’hypsodontie peut légèrement varier avec l’âge individuel et l’usure dentaire (Chaline 1972). Les spécimens de Ben Kérat présentent de plus une taille particulièrement importante, se situant dans la variabilité haute de E. barbarus, voire au-delà (Table 5; Fig. 12), les m2 et m3 sont peu réduites, et les molaires inférieures sont relativement larges (Table 6). Ainsi, les restes de Ben Kérat se rapprochent davantage de E. barbarus, malgré un indice d’hypsodontie plus modéré et une taille un peu plus importante. Devant la faible quantité de matériel et son mauvais état de préservation, nous préférons rester prudents et limiter l’attribution à E. cf. barbarus. Les différences observées pourraient aller dans le sens de l’hypothèse de «différentiations géographiques» liées à des conditions climatiques locales particulières déjà évoquée par Jaeger (1988) pour E. zimae par exemple., Published as part of Stoetzel, Emmanuelle & Pickford, Martin, 2022, Étude d'un assemblage original de microvertébrés du Pléistocène moyen du nord-est de l'Algérie (Ben Kérat, Oued Zenati) et description de deux nouveaux muridés, pp. 237-263 in Geodiversitas 44 (8) on page 251, DOI: 10.5252/geodiversitas2022v44a8, http://zenodo.org/record/6245987, {"references":["JAEGER J. J. 1988. - Origine et evolution du genre Ellobius (Mammalia, Rodentia) en Afrique nord-occidentale. Folia Quaternaria 57: 3 - 50.","MEIN P. & PICKFORD M. 1992. - Gisements karstiques pleistocenes au Djebel Ressas, Tunisie. Comptes Rendus de l'Academie des Sciences de Paris, Serie II 315: 247 - 253. https: // gallica. bnf. fr / ark: / 12148 / bpt 6 k 6304735 s / f 261. item","SAHNOUNI M., PARES J., DUVAL M., VAN DER MADE J., HARICHANE Z., PEREZ- GONZALEZ A., ABDESSADOK S., ARNOLD L., CACERES I., KANDI N., CHELLI- CHEHEB R., BOULAGHRAIF K., AGUSTI J., SAIDANI N. & MOUHOUBI Y. 2018 b. - Early Homo adaptive and behavioral patterns in North Africa: Perspectives from Ain Hanech and Tighennif (formerly Ternifine) sites in northern Algeria. Communication orale, 15 th PANAF Congress, 10 - 14 September 2018, Rabat, Morocco. Abstract Book, p. 5.","PICKFORD M. 2020. - The Fossil Suidae (Mammalia, Artiodactyla) from Ternifine (Tighenif) Algeria. Munchner Geowissenschaftliche Abhandlungen (A) 50: 1 - 66.","GERAADS D. 1994. - Rongeurs et Lagomorphes du Pleistocene moyen de la \" Grotte des Rhinoceros \", carriere Oulad Hamida 1, a Casablanca, Maroc. Neues Jahrbuch fur Geologie und Paleontologie, Abh 191 (2): 147 - 172.","GERAADS D. 2016 a. - La faune de vertebres du Pleistocene moyen de la Grotte des Rhinoceros, Casablanca, Maroc: 4 - Rodentia et Lagomorpha, in RAYNAL J. P. & MOHIB A. (eds), Prehistoire de Casablanca. 1 - La Grotte des Rhinoceros (fouilles 1991 et 1996). Villes et Sites Archeologiques du Maroc, Vol. VI, Rabat: 95 - 104.","CHALINE J. 1972. - Les rongeurs du Pleistocene moyen et superieur de France (systematique - biostratigraphie - paleoclimatologie). Cahiers de Paleontologie, Editions du Centre national de la Recherche scientifique, Paris, 410 p."]}

80. Alai Mole Vole Ellobius alaicus French: Campagnol des Alai / German: Alai-Mull-Lemmming / Spanish: Topillo topo de Alay Taxonomy. Ellobius alaicus Vorontsov et al., 1969, between Sary-Tashem and Bardabo, Alai Valley, S Kyrgyzstan. Ellobius alaicus is in the subgenus Ellobius. It is recognized as a species and distinct from E. tancrei based on chromosomal differences. Karyotype is 2n = 54 in tancre: and 2n = 52 in alaicus. Monotypic. Distribution. Alai Mts in S Kyrgyzstan. Descriptive notes. Head-body 110-132 mm, tail 8-17 mm; weight 42-6 63-8 g. The Alai Mole Vole is nearly identical in external morphology and craniodental structure to the Eastern Mole Vole (E. tancrei). Pelage is buff on back and gray along flanks and on belly. In some individuals, buff is shaded yellow, but in others, it is hued gray. Forehead is dark brown and does not contrast sharply with back. Cheeks are buff in some individuals. Incisors are strictly proodont. Molars are rooted, with no cement in reentrant loops. Habitat. Alpine pastures (type locality at an elevation of 3300 m). Food and Feeding. No information. Breeding. No information. Activity patterns. No information. Movements, Home range and Social organization. No information. Status and Conservation. Classified as Data Deficient on The IUCN Red List. There is a shortage of information on distribution, population size, trends, and threats to the Alai Mole Vole. Entire area of distribution is only ¢.2600 km?. Bibliography. Shenbrot & Krasnov (2005), Vorontsov & Yakimenko (1984), Vorontsov et al. (1969)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Cricetidae, pp. 204-535 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on page 314, DOI: 10.5281/zenodo.6707142

78. Northern Mole Vole Ellobius talpinus French: Campagnol taupe / German: Nordlicher Mull-Lemming / Spanish: Topillo topo septentrional Taxonomy. Mus talpinus Pallas, 1770, between Kuibyshev (= Samara) and Kostytchi, W bank of Volga River, Russia. Ellobius talpinus is in the subgenus Ellobius. In the past, E. talpinus also included E. tancrei. Taxonomic distinction between these two species was first retrieved from chromosomal evidence and subsequently confirmed by molecular data. They presumably diverged during Quaternary climatic cycles about one million years ago. Four subspecies recognized. Subspecies and Distribution. E.t.talpinusPallas,1770—SEEuropeanRussia,SWSiberia,andpossiblyNKazakhstan. E.t.rufescensEversmann,1850—WKazakhstan(betweenlowerUralandEmbarivers,andprobablydesertsWofAralSeaincludingNWUzbekistan). E.t.tanaiticusZubko,1940—UkraineandSEuropeanRussiaasfarEasNWCaspianshoreandthelowerVolgaRiver. E. t. transcaspiae Thomas, 1912 — Turkmenistan and NE Iran. Descriptive notes. Head-body 84-115 mm, tail 8-21 mm; weight 24-56 g. Body of the Northern Mole Vole is cylindrical with large, blunt head, short neck, and heavy chest. Nose pad is large, ears vestigial, and eyes are minute (3-2 mm in diameter). Tail is shorter than hindfoot, densely covered with stiff hairs. Front feet are heavier than hindfeet. Thumb is not reduced to such a degree as in most rodent species. Claws are strong but short. Females have eight nipples: two pairs of abdominal and two pairs of inguinal. Furis velvety, and coloris extremely variable. Belly is normally lighter and grayish. Head is darker than back, frequently black between nose and ears. Lightest individuals have sandy brown back, light gray belly, and brown head. In contrast, blackish brown individuals have black heads or are completely black; they can have rusty blotches on head and rusty front legs. In most populations of Northern Mole Voles, back is either grayish brown, brown, fawn, or rusty brown; belly is gray, occasionally washed buff; and head is dark brown or black. Skull is fossorial, relatively wider than species in subgenus Afganomys; rostrum is shorter; incisive foramina are less reduced; interorbital region is flat and wide; and supratemporal ridges neverfuse into crest. Mandible is powerful, with prominent alveolar process. Incisors are long and markedly fossorial, with white front surfaces. Molars are rooted, with simple pattern of broadly confluent dental fields and without cement in reentrant loops. Habitat. Forest steppe, steppe, and semi-deserts on chernozem (black-colored, humusrich) soil, clay, sand, and even gravelly or stony substrate. Northern Mole Voles are sensitive to freezing of soil during winter in northern borders of the distribution, and deep snow cover can be crucial for winter survival. During dry summers, they migrate from steppic habitats into birch (Betula, Betulaceae) stands. Food and Feeding. Northern Mole Voles consume up to 59-67 g of food/ day. Diet consists of underground parts (i.e. bulbs ofiris and garlic) and grass roots. Feeding tunnels are 15-20 cm deep in Turkmenistan. Feeding on the ground’s surface becomes more frequent in autumn and during reproduction. Food is not cached. Breeding. Reproduction starts in late winter and early spring and peaks in February— March. Females have 1-2 litters/year, but some authors have reported 3-4 litters/year. Mean litter sizes are 3-1-4-9 and vary among populations; overall mean for the species is four young (range 1-7). Gestation is 30 days. Newborns weight 3 g and are naked and blind. Eyes open at 21-28 days old, and young start eating solid food at 20-25 days old. Juveniles enter the population in April-May, rarely in March or June; they weight c.40 g at two months old. Females may reproduce during first year but more frequently are sexually mature in their second year. Life expectancy is two years. Activity patterns. Northern Mole Voles are active day and night, with peaks in morning and evening. Activity is reduced in winter and during summer droughts, and individuals can undergo lethargic periods. Northern Mole Voles are strictly fossorial. They rarely emerge from burrows and only at night. Digging activity goes on yearround, with peaks in spring and autumn. Speed of digging through soil averages 10-2 cm/minute. Digging is performed by two or more individuals. The first one digs with its incisors, and helpers behind push excavated dirt to the ground’s surface; positions are rotated. Amount of soil excavated annually is estimated at 2-3 m’/family group. Entrances to a burrow system are sealed with elliptically shaped mounds of soil, 13-68 cm long, 10-38 cm wide, and 2-10 cm high. There are 20-35 heaps/ family group, and excavated soil can cover 20-23% of the ground’s surface. Nest chamberis 11 x 11 x 9 cm and 40-65 cm deep in Turkmenistan but deeper (1-4 m) in the north. Old nest chambers are plugged with soil. In hot days and after heavy rains, Northern Mole Voles unplug burrows for ventilation. Tunnels are 5-6 cm in diameter (mean 5-2 cm). Movements, Home range and Social organization. The Northern Mole Vole is colonial, with family groups spaced 200-300 m apart. Burrow systems are dug out and maintained by a family of up to ten individuals. Only one female is reproductively active in a family. In Turkmenistan, postnatal dispersion is in March—-May. Young move up to 800 m away from natal burrows. Status and Conservation. Classified as Least Concern on The IUCN Red List. Populations of Northern Mole Voles on western edge of the distribution are declining. Entire population in Ukraine is believed to be 2000-3000 individuals, and it is extirpated on lower parts of the Dnieper River. Bibliography. Bogdanov et al. (2015), Evdokimov (2001), Gromov & Erbajeva (1995), Korobchenko & Kondratenko (2009), Marinina (2005b), Popov (1960), Shenbrot & Krasnov (2005), Shlyakhtin et al. (2009), Vorontsov & Yakimenko (1984)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Cricetidae, pp. 204-535 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on pages 313-314, DOI: 10.5281/zenodo.6707142

81. Southern Mole Vole Ellobius fuscocapillus French: Campagnol du Baloutchistan / German: Stdlicher Mull-Lemming / Spanish: Topillo topo meridional Other common names: Afghan Mole Vole, Quetta Mole Vole Taxonomy. Georychus fuscocapillus Blyth, 1843, Quetta, Baluchistan Region, Pakistan. Ellobius fuscocapillus is in the subgenus Af ganomys. Monotypic. Distribution. NE & SE Iran, W, S & E Turkmenistan, Afghanistan, and very marginally in W Pakistan (Balochistan). Descriptive notes. Head-body 98-141 mm (males) and 100-145 mm (females), tail 9:-1-19 mm (males) and 9-20 mm (females); weight 34-61 g (males) and 37-101 g (females). Females are slightly larger. The Southern Mole Vole is very similar to the Transcaucasian Mole Vole (FE. lutescens), but on average,it is larger and lighter. It is the largest species of Ellobius. Vestigial ear is 2-3 mm high. Fur is short, soft, and velvety, light yellowish brown to bufty gray on back and light gray on sides and belly. Some individuals are rustier, and others are pale yellow. Behind nasal pad and on forehead, pelage is shorter and decidedly darker, contrasting back and bright buff cheeks. This area is dark brown to blackish brown, lighter in winter and darker in summer. Subadults are lighter. Tail is brown, darker dorsally, and densely covered with hairs. Skull shows modifications for fossorial life. Zygomatic arches are widely expanded, sagittal and lambda crests are present in adult individuals, and incisive foramen is very short. Interparietal bone is missing. Mandible is powerful, with prominent alveolar process. Incisors are white, long, and markedly fossorial. They are exposed in front of the mouth even when lips are closed. Molars are rooted, with simple pattern of broadly confluent dentalfields and without cement in reentrant loops. Habitat. Open habitats with at least some grassy vegetation, frequently shaded with Ephedra (Ephedraceae) shrubs and junipers (Cupressaceae), at elevations of 300-2600 m. The Southern Mole Vole can tolerate gravelly or stony substrate but avoids flooded areas, steep rocky slopes, and moving sands. Valley bottoms and borders of cultivation with deep soil (loess or gravelly-clay) for digging are preferred. Food and Feeding. The Southern Mole Vole digs tunnels to obtain plants for feeding. In Balochistan, diet largely consists of pulpy tubers of foxtail lilies (Eremurus persicus, Asphodelaceae), garlic (Allium, Amaryllidaceae), and tulip (7 Tulipa, Liliaceae), but rhizomes of grasses are important in Turkmenistan. Food is cached, and stores are 25-30 g. Feeding on the ground’s surface is very exceptional. Breeding. In Turkmenistan, reproduction peaks in April-May. Females have 2-3 embryos, although counts up to five have been reported. Litters born in captivity had 1-4 young (mean 2-6). Activity patterns. The Southern Mole Vole is primarily crepuscular and nocturnal. It is fossorial, spending most ofits time underground. During digging, it braces hindlegs against the wall of the tunnel and pushes forward with its blunt muzzle. Proodont incisors act as a drill to loosen soil that is pushed back beneath belly by forefeet. Soil excavated from burrows is pressed to the ground’s surface in mounds. Peak digging activity is April-May. In hard soil, the Southern Mole Vole can dig deep enough to conceal its body in 15-19 minutes. Velvety fur minimizes friction in narrow corridors and enables backward movements with equal effectiveness as forward movements. Tunnels are 4.4-6 cm in diameter. Main tunnels are 15-25 cm deep, and branching feeding tunnels are short and shallow (6-10 cm deep). Nests are constructed at greater depths (35-115 cm), frequently in wet soil. Caches are 20-50 cm deep. Southern Mole Voles regulate temperature and humidity in burrows by plugging and unplugging entrances—activities that occur any time, including during the day. Movements, Home range and Social organization. Families of Southern Mole Voles occupy 25-80 m? that are 70-1500 m apart. Status and Conservation. Classified as Least Concern on The IUCN Red List. Area of occurrence of the Southern Mole Vole is ¢.500,000 km?. Bibliography. Gharkheloo & Kivang (2003), Marinina (2005a), Niethammer (1970), Roberts (1997), Shenbrot & Krasnov (2005), Yakimenko & Marochkina (2016)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Cricetidae, pp. 204-535 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on pages 314-315, DOI: 10.5281/zenodo.6707142

82. Transcaucasian Mole Vole Ellobius lutescens French: Campagnol fauve / German: Transkaukasischer Mull-Lemming / Spanish: Topillo topo del Caucaso Other common names: Mountain Mole Vole Taxonomy. Ellobius lutescens Thomas, 1897, “ Van, [Turkey], alt. 5000 feet [= 1524 m].” Ellobius lutescensis in the subgenus Afganomys. Monotypic. Distribution. E Turkey (E Anatolia), SW Azerbaijan (Nakhchivan), Armenia, and NW Iran. Descriptive notes. Head-body 100-140 mm, tail 7-19 mm; weight 58-84 g. The Transcaucasian Mole Vole is small, with rounded head and short, hairy tail with long terminal pencil. Nose pad is large, eyes are minute, and ears are reduced to skinfolds around ear canals. Whiskers are directed forward, stiff and up to 14-5 mm long. Feet are large and robust, and their lateral borders are edged by rows of stiff hairs. Claws are strong and short. Females have two pairs of pectoral and one pair of inguinal nipples (six nipples in total), but some sources report two inguinal pairs (eight nipples in total). Fur is short, soft, and dense, slate buffy throughout but lighter on belly. Pelt is darkest around eyes and on front and behind nose pad. Some individuals have white blotches in different parts, frequently on undersides. Young voles are lighter than adults. Skull is short, wide, and deep, with bowed dorsal surface. Interorbital region is wide and without sagittal crest. Diastema is long, and incisive foramina are small and shifted posteriorly. Mandible is heavy, with massive ramus and protruding alveolar process. Incisors are long and strongly proodont. Molars are rooted, with simple pattern of enamel loops and without cement. Dental fields are broadly confluent. Habitat. Dry grasslands and arable land at elevations of 600-3000 m. Food and Feeding. Diet of the Transcaucasian Mole Vole consists primarily of bulbs, tubers, and roots of cranesbill (Geranium, Geraniaceae), earthnut (Bunium, Apiaceae), and garlic (Allium, Amaryllidaceae). Potatoes, carrots, and onions are consumed in cultivated areas, frequently at depths of 10-12 cm. Food is stored in expanded nest chambers. Daily consumption is 0-74 g per gram of body mass. Breeding. Female Transcaucasian Mole Voles are sexually mature at 16-17 weeks old. Pregnant females and juveniles were collected in March—October, but reproduction evidently peaks in spring. Gestation is 21 days, and shortest interval between two births in captivity was 28 days. Females have 2—4 embryos, and litter sizes in captivity were 2-3 young (mean 2-6). Young are born naked, with closed eyes and average weight of 3-4 g. They have one pair each of upper and lower incisors that facilitate clinging onto nipples. These incisors are lost at 19-21 days old. Young start feeding on solid food at 23-24 days old, while they are still suckling, and they are weaned at 33-35 days old. Eyes open at 26-28 days old. After three months, adult size is reached, but incisors are still shorter and pelage is brighter than those of adults. Activity patterns. Transcaucasian Mole Voles are active day and night, with peaks at dusk and dawn. Most activity is restricted to underground galleries. They dig with proodont incisors and push loosen soil beneath and out of tunnels. Mounds of excavated soil are 20-50 cm in diameter and 9-14 cm high. Tunnels with diameters of 6-9 cm are 16-48 cm below the ground’s surface. Emergency tunnels descend 54— 90 cm. Nest chamberis lined with dry grass and other soft material (e.g. plastic and textile), and it can be expanded to act as storing chamber. Movements, Home range and Social organization. In eastern Turkey, home range of a family of Transcaucasian Mole Voles was 108 m®. In captivity, parents coexisted peacefully with up to three litters. Status and Conservation. Classified as Least Concern on The IUCN Red List. Overall distribution of the Transcaucasian Mole Vole is ¢.310,000 km?. Bibliography. Coskun & Ulutlrk (2003), Gharkheloo & Kivang (2003), Kaya & Coskun (2015), Krystufek & Vohralik (2005), Shenbrot & Krasnov (2005)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Cricetidae, pp. 204-535 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on page 315, DOI: 10.5281/zenodo.6707142

The subterranean mole vole, Ellobius tancrei, with a specific variability in autosomes (2n = 31–54) and unusual sex chromosomes (XX in males and females), represents an amazing model for studying the role of chromosome changes in speciation. New materials from the upper reaches of the Surkhob River in the Pamiro-Alay mountains resulted in the discovery of a new form with 2n = 30. The application of Zoo-FISH and G-banding methods allowed the detection of 13 pairs of autosomes as Robertsonian metacentrics originated after fusions of acrocentrics of an assumed ancestral karyotype of E. tancrei with 2n = 54. The sex chromosomes (XX, in both sexes) and one pair of acrocentric autosomes are the only acrocentrics in this karyotype, and the set with 2n = 30 possesses the lowest possible chromosome number among populations of E. tancrei.

The synaptonemal complex (SC) surface-spreading technique was used to visualize the process of chromosome synapsis in spermatocytes and oocytes of E. talpinus Pallas, 1770, a species with the XX sex chromosome system in both males and females. We used electron microscopy and immunofluorescent localization of synaptonemal complex protein (SCP3) and centromeric proteins to analyze the structure and behaviour of synaptonemal complexes in prophase I of meiosis, aiming to reveal signs of meiotic sexual dimorphism in this species. We present evidence of considerable differences in the structure and behaviour of the axial structures of sex bivalents in male and female meiosis, despite the isomorphic G- and C-banding patterns of mitotic sex chromosomes. During meiotic prophase I, the sex bivalent in females behaved as autosomal bivalents, but it was not involved in the formation of the bouquet configuration or it was the first to leave it. The XX chromosomes of males formed closed sex bivalents. Only short tracts of SC were formed at both ends of the sex bivalent, while large middle segments of the lateral elements remained unpaired. The male sex chromosomes also formed characteristic “sex bodies”. In fact, electron microscopy revealed dense nucleolus-like bodies associated with unpaired parts of the axial elements. These regions of the sex chromosomes were poorly immunostained, because the distribution of SCP3 had a peculiar powder-like pattern, but SCP3 was not associated with the nucleolus-like bodies. We also revealed signs of sexual dimorphism in the dynamics of formation and destruction of autosomal SCs. In males, the total SC length was shorter than in females. The chromosome bouquet configuration was preserved up to the stage of early pachytene in females. The bouquet configuration in males was not expressed. At late pachytene, gaps were revealed in the structure of autosomal SCs in spermatocytes immunostained with antibodies to SCP3. The pattern of distribution of these gaps was comparable with the G-banding patterns of mitotic chromosomes.

Ellobius tancrei Blasius, 1884. Zool. Anz., 7:197. TYPE LOCALITY: Kazakhstan, Zaissan Lake Valley, Kendyrlik (= Przevalskoie). DISTRIBUTION: From NE Turkmenistan (Yakimenko and Lyapunova, 1986) and Uzbekistan east through E Kazakhstan to E China (Xinjiang and Nei Mongolia) and Mongol. SYNONYMS: coenosus, fusciceps, fuscipes, kastschenkoi, ognevi, ursulus. COMMENTS: Included in E. talpinus by Corbet (1978c) but now regarded as a distinct species whose geographic range is allopatric to that of E. talpinus (see references in Corbet, 1984; Pavlinov and Rossolimo, 1987; Yakimenko and Lyapunova, 1986). Chromosomal contrasts with E. talpinus recorded by Yakimenko and Lyapunova (1986), comparisons with E. alaicus reported by Lyapunova et al. (1990)., Published as part of Guy G. Musser & Michael D. Carleton, 1993, Order Rodentia - Family Muridae, pp. 501-755 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 513, DOI: 10.5281/zenodo.7353098, {"references":["Yakimenko, L. V., and E. A. Lyapunova. 1986. [Cytogenetic corroboration of belonging of northern mole vole from Turkmenia to Ellobius talpinus s. str.] Zoologicheskii Zhurnal, 65: 946 - 949 (in Russian).","Corbet, G. B. 1978 c. The mammals of the Palaearctic region: A taxonomic review. British Museum (Natural History), London, 314 pp.","Corbet, G. B. 1984. The mammals of the Palearctic region: A taxonomic review. Supplement. British Museum (Natural History), London, 45 pp.","Pavlinov, I. Ya., and O. L. Rossolimo. 1987. Sistematika mlekopitayushchikh SSSR [Systematics of the mammals of the USSR.]. Moscow University Press, Moscow, 282 pp. (in Russian)."]}

Ellobius alaicus Vorontsov et al., 1969. In Vorontsov (ed.), [The Mammals: Evolution, karyology, taxonomy, fauna], Novosibirsk, p. 127. TYPE LOCALITY: Kirghizia, Alai Valley, between Sary-Tashem and Bardabo, 3300 m. DISTRIBUTION: Recorded only from the Alai Mtns, S Kirghizia. COMMENTS: Subgenus Ellobius. Included, with question, in E. talpinus by Corbet (1978c). Chromosomal data and breeding results reveal that alaicus does not belong in E. talpinus and is reproductively isolated from E. tancrei, with which it is parapatric (Corbet, 1984; Lyapunova et al., 1990, and references therein). Closely related to E. tancrei and provisionally placed in that species by Pavlinov and Rossolimo (1987)., Published as part of Guy G. Musser & Michael D. Carleton, 1993, Order Rodentia - Family Muridae, pp. 501-755 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 512, DOI: 10.5281/zenodo.7353098, {"references":["Corbet, G. B. 1978 c. The mammals of the Palaearctic region: A taxonomic review. British Museum (Natural History), London, 314 pp.","Corbet, G. B. 1984. The mammals of the Palearctic region: A taxonomic review. Supplement. British Museum (Natural History), London, 45 pp.","Pavlinov, I. Ya., and O. L. Rossolimo. 1987. Sistematika mlekopitayushchikh SSSR [Systematics of the mammals of the USSR.]. Moscow University Press, Moscow, 282 pp. (in Russian)."]}

Ellobius fuscocapillus Blyth, 1843. J. Asiat. Soc. Bengal, 11:887. TYPE LOCALITY: Pakistan, Baluchistan Region, Quetta Div., Quetta. DISTRIBUTION: E Iran, Afghanistan, W Pakistan, and S Turkmenistan in the Kopet Dag Mtns. SYNONYMS: farsistani, intermedius. COMMENTS: Subgenus Afganomys. Chromosomal data presented by Vorontsov et al. (1980) and Lyapunova et al. (1980) in context of assessing specific differences within the genus and variation in sex chromosomes. Morphology described in detail by Hinton (1926), in context of surveying interrelationships among arvicoline genera., Published as part of Guy G. Musser & Michael D. Carleton, 1993, Order Rodentia - Family Muridae, pp. 501-755 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 512, DOI: 10.5281/zenodo.7353098, {"references":["Vorontsov, N. N., E. A. Lyapunova, Yu. M. Borisov, and V. E. Dovgal. 1980. Variability of sex chromosomes in mammals. Genetica, 52 / 53: 361 - 372.","Lyapunova, E. A., N. N. Vorontsov, K. V. Korobitsyna, E. Yu. Ivanitskaya, Yu. M. Borisov, L. V. Yakimenko, and V. Ye. Dovgal. 1980. A Robertsonian fan in Ellobius talpinus. Genetica, 52 / 53: 239 - 247."]}

Ellobius lutescens Thomas, 1897. Ann. Mag. Nat. Hist., ser. 6, 20:308. TYPE LOCALITY: Turkey, Kurdistan, Van. DISTRIBUTION: From S Caucasus Mtns south through E Turkey and NW Iran. COMMENTS: Subgenus Afganomys. Treated by Corbet (1978c) as a synonym of E. fuscocapillus but now shown to be a distinct species (Corbet, 1984; Vorontsov et al., 1980, and references therein), as earlier listed by Ellerman and Morrison-Scott (1951). This is a species with a low diploid number (17) and an unusual mode of sex determination, which has been the subject of the many reports reviewed by Zima and Krâl (1984a), as well as current inquiry (Vogel et al., 1988, and references therein)., Published as part of Guy G. Musser & Michael D. Carleton, 1993, Order Rodentia - Family Muridae, pp. 501-755 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 512, DOI: 10.5281/zenodo.7353098, {"references":["Corbet, G. B. 1978 c. The mammals of the Palaearctic region: A taxonomic review. British Museum (Natural History), London, 314 pp.","Corbet, G. B. 1984. The mammals of the Palearctic region: A taxonomic review. Supplement. British Museum (Natural History), London, 45 pp.","Vorontsov, N. N., E. A. Lyapunova, Yu. M. Borisov, and V. E. Dovgal. 1980. Variability of sex chromosomes in mammals. Genetica, 52 / 53: 361 - 372.","Ellerman, J. R., and T. C. S. Morrison-Scott. 1951. Checklist of Palaearctic and Indian mammals 1758 to 1946. Trustees of the British Museum (Natural History), London, 810 pp.","Zima, J., and B. Kral. 1984 a. Karyotypes of European mammals. II. Acta Scientiarum Naturalium, Academiae Scientarium Bohemoslovacae (Brno), 18 (8): 1 - 62.","Vogel, W., P. Steinbach, M. Djalali, K. Mehnert, S. Ali, and J. T. Epplen. 1988. Chromosome 9 of Ellobius lutescens is the X chromosome. Chromosoma, 96: 112 - 118."]}

Ellobius talpinus (Pallas, 1770). Nova Comm. Acad. Sci. Petropoli, 14, 1:568. TYPE LOCALITY: Russia, W Bank of Volga River, between Kuibyshev (= Samara) and Kostychi. DISTRIBUTION: Steppes from S Ukraine and Crimea east through Kazakhstan to N of Balkhash Lake, and in Turkmenistan (see Pavlinov and Rossolimo, 1987; Yakimenko and Lyapunova, 1986). SYNONYMS: ciscaucasicus, murinus, rufescens, tanaiticus, transcaspiae. COMMENTS: Subgenus Ellobius. Chromosomal polymorphism among samples from the Pamir-Alai Mtns was analyzed by Lyapunova et al. (1980); other chromosomal data summarized and reviewed by Zima and Krâl (1984a). Cytological identification of Turkmenian populations as E. talpinus and comparison with T. tancrei provided by Yakimenko and Lyapunova (1986)., Published as part of Guy G. Musser & Michael D. Carleton, 1993, Order Rodentia - Family Muridae, pp. 501-755 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 512, DOI: 10.5281/zenodo.7353098, {"references":["Pavlinov, I. Ya., and O. L. Rossolimo. 1987. Sistematika mlekopitayushchikh SSSR [Systematics of the mammals of the USSR.]. Moscow University Press, Moscow, 282 pp. (in Russian).","Yakimenko, L. V., and E. A. Lyapunova. 1986. [Cytogenetic corroboration of belonging of northern mole vole from Turkmenia to Ellobius talpinus s. str.] Zoologicheskii Zhurnal, 65: 946 - 949 (in Russian).","Lyapunova, E. A., N. N. Vorontsov, K. V. Korobitsyna, E. Yu. Ivanitskaya, Yu. M. Borisov, L. V. Yakimenko, and V. Ye. Dovgal. 1980. A Robertsonian fan in Ellobius talpinus. Genetica, 52 / 53: 239 - 247.","Zima, J., and B. Kral. 1984 a. Karyotypes of European mammals. II. Acta Scientiarum Naturalium, Academiae Scientarium Bohemoslovacae (Brno), 18 (8): 1 - 62."]}

Ellobius Fischer, 1814. Zoognosia, 3:72. TYPE SPECIES: Mus talpinus Pallas, 1770. SYNONYMS: Afganomys, Afghanomys, Chthonergus, Lemmomys, Myospalax (of Blyth, 1846, not Laxmann, 1769, or Hermann, 1783). COMMENTS: Gromov and Polyakov (1977) excluded Ellobius from arvicolines, and Pavlinov and Rossolimo (1987) viewed the Ellobiini as Cricetidae incertae sedis, questioning whether it belonged in Arvicolinae or Cricetinae. Most workers, however, have recognized Ellobius, albeit highly specialized and unusual, as the only extant member of the tribe Ellobiini of Arvicolinae (Corbet, 1978c; Hooper and Hart, 1962; Kretzoi, 1969; Topachevskii and Rekovets, 1982). Topachevskii and Rekovets (1982) interpreted relationships and morhological trends in diversification within the genus from the late Pliocene to Recent. The present range of Ellobius is only part of a wider distribution that once embraced Israel and North Africa in the middle to late Pleistocene (Jaeger, 1988). Ultrastructure, meiotic behavior, and evolution of sex chromosomes in the genus are discussed by Kolomiets et al. (1991). Two subgenera are recognized, Ellobius and Afganomys; Zagorodnyuk (1990) listed the latter as a genus., Published as part of Guy G. Musser & Michael D. Carleton, 1993, Order Rodentia - Family Muridae, pp. 501-755 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 512, DOI: 10.5281/zenodo.7353098, {"references":["Hermann, J. 1783. Tabula Affinitatum Anim. p. 115.","Gromov, I. M., and I. Ya. Polyakov. 1977. Fauna SSSR, Mlekopitayushchie, tom 3, vyp. 8 [Fauna of the USSR, vol. 3, pt. 8, Mammals]. Polevki [Voles (Microtinae)]. Nauka, Moscow-Leningrad, 504 pp. (in Russian).","Pavlinov, I. Ya., and O. L. Rossolimo. 1987. Sistematika mlekopitayushchikh SSSR [Systematics of the mammals of the USSR.]. Moscow University Press, Moscow, 282 pp. (in Russian).","Corbet, G. B. 1978 c. The mammals of the Palaearctic region: A taxonomic review. British Museum (Natural History), London, 314 pp.","Hooper, E. T., and B. S. Hart. 1962. A synopsis of Recent North American microtine rodents. Miscellaneous Publications, Museum of Zoology, University of Michigan, 120: 1 - 68.","Kretzoi, M. 1969. Skizze einer Arvicoliden-Phylogenie. Vertebrata Hungarica (Budapest), 11: 155 - 193.","Topachevskii, V. A., and L. I. Rekovets. 1982. Novye materialy k sistematike i evolyutsii slepushonok nominativnovo podrod roda Ellobius (Rodentia, Cricetidae) [New material for systematic and evolution of the mole vole, nominative subgenus and genus Ellobius (Rodentia, Cricetidae)]. Vestnik Zoologii, 5: 47 - 54 (in Russian).","Kolomiets, O. L., N. N. Vorontsov, E. A. Lyapunova, and T. F. Mazurova. 1991. Ultrastructure, meiotic behavior, and evolution of sex chromosomes of the genus Ellobius. Genetica, 84: 179 - 189.","Zagorodnyuk, I. V. 1990. Kariotipicheskaya izmenchivost' i sistematika serykh polevok (Rodentia, Arvicolini). Soobshchenie 1. Vidovoi sostav i khromosomnye chisla [Karyotypic variability and systematics of the gray voles (Rodentia, Arvicolini). Communication 1. Species composition and chromosomal numbers]. Vestnik Zoologii, 2: 26 - 37 (in Russian)."]}

Ellobius lutescens Thomas, 1897. Ann. Mag. Nat. Hist., ser. 6, 20:308. TYPE LOCALITY: Turkey, Kurdistan, Van. DISTRIBUTION: S. Caucasus Mtns. (U.S. S.R.), south through E. Turkey and W. Iran; perhaps Iraq. COMMENT: Corbet, 1978:117, and others considered lutescens a synonym of fuscocapillus; but Lyapunova and Vorontsov, 1978, Genetika, 14: 2012-2024, and Lyapunova et al., 1980:293-247, in Vorontsov and Van Brink, eds., Animal genetics and evolution, Junk, The Hague, 383 pp., provided evidence of specific distinctness, ISIS NUMBER: 5301410008075002001., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Rodentia (Part 4), pp. 477-504 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 485, DOI: 10.5281/zenodo.7353034, {"references":["Vorontsov, N. N., E. A. Lyapunova, E. Yu. Ivanitskaya, C. F. Nadler, B. Kral, A. I. Kozlovskii, and R. S. Hoffmann. 1978. [Variability of mammalian sex chromosomes. I. Geographic variability of the structure of the Y chromosomes in red-backed voles of the genus Clethrionomys (Rodentia, Microtinae).] Genetika, 14: 1432 - 1446.","Lyapunova, E. A., N. N. Vorontsov, K. V. Korobitsyna, E. Yu. Ivanitskaya, Yu. M. Borisov, L. V. Yakimenko, and V. Ye. Dovgal. 1980. A Robertsonian fan in Ellobius talpinus. Genetica, 52 / 53: 239 - 247."]}

Ellobius fuscocapillus Blyth, 1843. J. Asiat. Soc. Bengal, 11:887. TYPE LOCALITY: Pakistan, Baluchistan Reg., Quetta Div., Quetta. DISTRIBUTION: Baluchistan and Afghanistan through Iran and S. Turkmenistan to Kurdistan. COMMENT: Formerly included lutescens; see Corbet, 1978:117, but also see Lyapunova and Vorontsov, 1978, Genetika, 14:2012-2024, and Lyapunova et al., 1980:239-247, in Vorontsov and Van Brink, eds., Animal genetics and evolution, Junk, The Hague, 383 pp. ISIS NUMBER: 5301410008075001001., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Rodentia (Part 4), pp. 477-504 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 484, DOI: 10.5281/zenodo.7353034, {"references":["Vorontsov, N. N., E. A. Lyapunova, E. Yu. Ivanitskaya, C. F. Nadler, B. Kral, A. I. Kozlovskii, and R. S. Hoffmann. 1978. [Variability of mammalian sex chromosomes. I. Geographic variability of the structure of the Y chromosomes in red-backed voles of the genus Clethrionomys (Rodentia, Microtinae).] Genetika, 14: 1432 - 1446.","Lyapunova, E. A., N. N. Vorontsov, K. V. Korobitsyna, E. Yu. Ivanitskaya, Yu. M. Borisov, L. V. Yakimenko, and V. Ye. Dovgal. 1980. A Robertsonian fan in Ellobius talpinus. Genetica, 52 / 53: 239 - 247."]}

Ellobius Fischer, 1814. Zoognosia, 3:72. COMMENT: Placed with Prometheomys in subfamily Prometheomyinae by CAR. Gromov, 1972, Sbor. Tr. Zool. Mus. M.G.U., 13:8 -32, excluded Ellobius from Arvicolidae as a microtodontine cricetid. OLR follows Corbet, 1978:117, in recognizing only two species fuscocapillus and talpinus; but see Lyapunova and Vorontsov, 1978, Genetika, 14: 2012-2024, and Gromov and Baranova, 1981:151. See comment under Reithrodon physodes (Cricetidae), for discussion of Fischer, 1814, names. ISIS NUMBER: 5301410008075000000., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Rodentia (Part 4), pp. 477-504 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 484, DOI: 10.5281/zenodo.7353034, {"references":["Vorontsov, N. N., E. A. Lyapunova, E. Yu. Ivanitskaya, C. F. Nadler, B. Kral, A. I. Kozlovskii, and R. S. Hoffmann. 1978. [Variability of mammalian sex chromosomes. I. Geographic variability of the structure of the Y chromosomes in red-backed voles of the genus Clethrionomys (Rodentia, Microtinae).] Genetika, 14: 1432 - 1446.","Gromov, I. M., and G. I. Baranova (eds.). 1981. Katalog mlekopitayushchikh SSSR [Catalog of mammals of the USSR]. Nauka, Leningrad, 456 pp. (in Russian)."]}

Ellobius talpinus (Pallas, 1770). Nova Comm. Acad. Sci. Petrop., 14(2): 568. REVIEWED BY: M. Andera (MA). TYPE LOCALITY: U.S. S. R., R.S.F.S. R., Kuibyshevsk. Obi., W. Bank of Volga River, between Kuibyshev (=Samara) and Kostychi. DISTRIBUTION: Steppes from S. Ukraine and Crimea, east through Kazakhstan (U.S. S.R.) to Sinkiang and Inner Mongolia (China) and Mongolia, north to Sverdlovsk and south to N. Afghanistan, N.W. Iran and N. Pakistan. COMMENT: Includes tancrei, which was considered a distinct species by Gromov and Baranova, 1981: 152. Karyology reviewed by Lyapunova et al., 1980:239-247, in Vorontsov and Van Brink, eds., Animal genetics and evolution, Junk, The Hague. ISIS NUMBER: 5301410008075003001., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Rodentia (Part 4), pp. 477-504 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 485, DOI: 10.5281/zenodo.7353034, {"references":["Gromov, I. M., and G. I. Baranova (eds.). 1981. Katalog mlekopitayushchikh SSSR [Catalog of mammals of the USSR]. Nauka, Leningrad, 456 pp. (in Russian).","Lyapunova, E. A., N. N. Vorontsov, K. V. Korobitsyna, E. Yu. Ivanitskaya, Yu. M. Borisov, L. V. Yakimenko, and V. Ye. Dovgal. 1980. A Robertsonian fan in Ellobius talpinus. Genetica, 52 / 53: 239 - 247."]}

Ellobius alaicus Vorontsov et al., 1969. In [The Mammals: Evolution, karyology, taxonomy, fauna], Sib. Otd., Akad. Nauk., Novosibirsk, p. 127. TYPE LOCALITY: U.S. S. R., Kirgiz S.S. R., Oshsk. Obi., Alai Valley between Sary-Tash and Bardabo. DISTRIBUTION: Alai Valley and Alai Mtns. (Kirgizia, U.S. S.R.). COMMENT: Included in talpinus by Corbet, 1978:117, but Lyapunova and Vorontsov, 1978, Genetika, 14:2012-2024, provided evidence of specific distinctness, although they agreed it is closely related., Published as part of James H. Honacki, Kenneth E. Kinman & James W. Koeppl, 1982, Order Rodentia (Part 4), pp. 477-504 in Mammal Species of the World (1 st Edition), Lawrence, Kansas, USA :Alien Press, Inc. & The Association of Systematics Collections on page 484, DOI: 10.5281/zenodo.7353034, {"references":["Vorontsov, N. N., E. A. Lyapunova, E. Yu. Ivanitskaya, C. F. Nadler, B. Kral, A. I. Kozlovskii, and R. S. Hoffmann. 1978. [Variability of mammalian sex chromosomes. I. Geographic variability of the structure of the Y chromosomes in red-backed voles of the genus Clethrionomys (Rodentia, Microtinae).] Genetika, 14: 1432 - 1446."]}