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ABSTTRACT: -Using the methods of geometric morphometrics, we have revealed the phenomenon of geographical variability of morphogenetic trajectories when doing side-by-side comparisons of mandibular shapes in individuals of different ages from three southern Trans-Ural populations of the mole vole (Ellobius talpinus Pall.) in the common morphospace.

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."]}

This study evaluates signatures of selection in the evolution of the mitochondrial DNA of voles, subfamily Arvicolinae, during the colonization of subterranean environments. The comparative sequence analysis of mitochondrial protein-coding genes of eight subterranean vole species (Prometheomys schaposchnikowi, three species of the genus Ellobius: Ellobius talpinus, Ellobius fuscocapillus and Ellobius lutescens, two species of the genus Terricola: Terricola subterraneus and Terricola daghestanicus, Lasiopodomys mandarinus, and Hyperacrius fertilis) and their closest aboveground relatives was applied using codon-substitution models. The highest number of selection signatures was detected in genes ATP8 and CYTB. The relaxation of selection was observed in most mitochondrial DNA protein-coding genes for subterranean species. The largest amount of relaxed genes is discovered in mole voles (genus Ellobius). The number of selection signatures was found to be independent of the evolutionary age of the lineage but fits the degree of specialization to the subterranean niche. The common trends of selective pressures were observed among the evolutionary ancient and highly specialized subterranean rodent families and phylogenetically young lineages of voles. It suggests that the signatures of adaptation in individual mitochondrial protein-coding genes associated with the colonization of the subterranean niche may appear within a rather short evolutionary timespan.

Evolutionary history and taxonomic position for cryptic species may be clarified by using molecular and cytogenetic methods. The subterranean rodent, the Alay mole voleEllobiusalaicusVorontsov et al., 1969 is one of three sibling species constituting the subgenus Ellobius Fischer, 1814, all of which lost the Y chromosome and obtained isomorphic XX sex chromosomes in both males and females.E.alaicusis evaluated by IUCN as a data deficient species because their distribution, biology, and genetics are almost unknown. We revealed specific karyotypic variability (2n = 52–48) inE.alaicusdue to different Robertsonian translocations (Rbs). Two variants of hybrids (2n = 53, different Rbs) withE.tancreiBlasius, 1884 were found at the Northern slopes of the Alay Ridge and in the Naryn district, Kyrgyzstan. We described the sudden change in chromosome numbers from 2n = 50 to 48 and specific karyotype structure for mole voles, which inhabit the entrance to the Alay Valley (Tajikistan), and revealed their affiliation asE.alaicusby cytochromeband fragments of nuclearXISTandRspo1genes sequencing. To date, it is possible to expand the range ofE.alaicusfrom the Alay Valley (South Kyrgyzstan) up to the Ferghana Ridge and the Naryn Basin, Tien Shan at the north-east and to the Pamir-Alay Mountains (Tajikistan) at the west. The closeness ofE.tancreiandE.alaicusis supported, whereas specific chromosome and molecular changes, as well as geographic distribution, verified the species status forE.alaicus. The case ofEllobiusspecies accented an unevenness in rates of chromosome and nucleotide changes along with morphological similarity, which is emblematic for cryptic species.

Cyclin-dependent kinases (CDKs) are crucial regulators of the eukaryotic cell cycle. The critical role of CDK2 in the progression of meiosis was demonstrated in a single mammalian species, the mouse. We used immunocytochemistry to study the localization of CDK2 during meiosis in seven rodent species that possess hetero- and homomorphic male sex chromosomes. To compare the distribution of CDK2 in XY and XX male sex chromosomes, we performed multi-round immunostaining of a number of marker proteins in meiotic chromosomes of the rat and subterranean mole voles. Antibodies to the following proteins were used: RAD51, a member of the double-stranded DNA break repair machinery; MLH1, a component of the DNA mismatch repair system; and SUN1, which is involved in the connection between the meiotic telomeres and nuclear envelope, alongside the synaptic protein SYCP3 and kinetochore marker CREST. Using an enhanced protocol, we were able to assess the distribution of as many as four separate proteins in the same meiotic cell. We showed that during prophase I, CDK2 localizes to telomeric and interstitial regions of autosomes in all species investigated (rat, vole, hamster, subterranean mole voles, and mole rats). In sex bivalents following synaptic specificity, the CDK2 signals were distributed in three different modes. In the XY bivalent in the rat and mole rat, we detected numerous CDK2 signals in asynaptic regions and a single CDK2 focus on synaptic segments, similar to the mouse sex chromosomes. In the mole voles, which have unique XX sex chromosomes in males, CDK2 signals were nevertheless distributed similarly to the rat XY sex chromosomes. In the vole, sex chromosomes did not synapse, but demonstrated CDK2 signals of varying intensity, similar to the rat X and Y chromosomes. In female mole voles, the XX bivalent had CDK2 pattern similar to autosomes of all species. In the hamster, CDK2 signals were revealed in telomeric regions in the short synaptic segment of the sex bivalent. We found that CDK2 signals colocalize with SUN1 and MLH1 signals in meiotic chromosomes in rats and mole voles, similar to the mouse. The difference in CDK2 manifestation at the prophase I sex chromosomes can be considered an example of the rapid chromosome evolution in mammals.

The subterranean voles of the genus Ellobius are species of subfamily Arvicolinae well adapted to underground life. In this paper, we report the assemblies of complete mitochondrial genomes for three mole voles from genus Ellobius – northern mole vole Ellobius talpinus (16,376 bp), transcaucasian mole vole E. lutescens (16,540 bp), and southern mole vole E. fuscocapillus (16,388 bp). Each of three mitogenomes encode for 12S and 16S rRNAs, 22 tRNAs, 13 protein-coding genes, and D-loop in the characteristic arrangement of subfamily Arvicolinae (Rodentia: Cricetidae). This study verifies the evolutionary status of subgenera Bramus and Ellobius within the genus Ellobius at the molecular level. The mitochondrial genome would be a significant supplement for the Ellobius genetic background. The three Ellobius species formed a monophyletic group with the high bootstrap value (100%) in all examinations.

International audience; Ellobius remains are common and often abundant in southeastern Europe, western and central Asia archaeological sites. A correct identification of species is crucial for our understanding of the evolution of species and communities through time, including biostratigraphic sequences to be established. This study applies geometric morphometric methods (GMM) to Ellobius first lower molars, with the objectives: 1) to discriminate modern species and explore morphological and size differences in reference samples; and 2) to identify fossil specimens recovered in archaeological sites, based on the aforementioned analysis. The reference dataset used in this paper includes specimens belonging to the three species that today occur in the southeastern Europe, western and central Asia: Ellobius fuscocapillus, E. lutescens and E. talpinus. The archaeological material comes from Late Pleistocene Iranian site of Kaldar Cave (Khorramabad valley, Lorestan Province, western Iran). Our study shows that the shape of the anterior cap and the arrangement of the following triangles allow discriminating the three studied extant Ellobius species. The shapes of E. fuscocapillus and E. lutescens m1 appear rather similar, whereas Ellobius talpinus is well separated from these two species. The total length and the anterior cap of m1 in E. fuscocapillus is greater than in Ellobius lutescens. The GMM analyses performed on the modern reference dataset allowed us to identify fossil specimens from Kaldar Cave as E. lutescens and some as E. fuscocapillus, and excluding E. talpinus.

Sex determination in mammals is usually provided by a pair of chromosomes, XX in females and XY in males. Mole voles of the genus Ellobius are exceptions to this rule. In Ellobius tancrei, both males and females have a pair of XX chromosomes that are indistinguishable from each other in somatic cells. Nevertheless, several studies on Ellobius have reported that the two X chromosomes may have a differential organization and behavior during male meiosis. It has not yet been demonstrated if these differences also appear in female meiosis. To test this hypothesis, we have performed a comparative study of chromosome synapsis, recombination, and histone modifications during male and female meiosis in E. tancrei. We observed that synapsis between the two X chromosomes is limited to the short distal (telomeric) regions of the chromosomes in males, leaving the central region completely unsynapsed. This uneven behavior of sex chromosomes during male meiosis is accompanied by structural modifications of one of the X chromosomes, whose axial element tends to appear fragmented, accumulates the heterochromatin mark H3K9me3, and is associated with a specific nuclear body that accumulates epigenetic marks and proteins such as SUMO-1 and centromeric proteins but excludes others such as H3K4me, ubiH2A, and γH2AX. Unexpectedly, sex chromosome synapsis is delayed in female meiosis, leaving the central region unsynapsed during early pachytene. This region accumulates γH2AX up to the stage in which synapsis is completed. However, there are no structural or epigenetic differences similar to those found in males in either of the two X chromosomes. Finally, we observed that recombination in the sex chromosomes is restricted in both sexes. In males, crossover-associated MLH1 foci are located exclusively in the distal regions, indicating incipient differentiation of one of the sex chromosomes into a neo-Y. Notably, in female meiosis, the central region of the X chromosome is also devoid of MLH1 foci, revealing a lack of recombination, possibly due to insufficient homology. Overall, these results reveal new clues about the origin and evolution of sex chromosomes.

Genome functioning in hybrids faces inconsistency. This mismatch is manifested clearly in meiosis during chromosome synapsis and recombination. Species with chromosomal variability can be a model for exploring genomic battles with high visibility due to the use of advanced immunocytochemical methods. We studied synaptonemal complexes (SC) and prophase I processes in 44-chromosome intraspecific (Ellobius tancrei &times, E. tancrei) and interspecific (Ellobius talpinus &times, E. tancrei) hybrid mole voles heterozygous for 10 Robertsonian translocations. The same pachytene failures were found for both types of hybrids. In the intraspecific hybrid, the chains were visible in the pachytene stage, then 10 closed SC trivalents formed in the late pachytene and diplotene stage. In the interspecific hybrid, as a rule, SC trivalents composed the SC chains and rarely could form closed configurations. Metacentrics involved with SC trivalents had stretched centromeres in interspecific hybrids. Linkage between neighboring SC trivalents was maintained by stretched centromeric regions of acrocentrics. This centromeric plasticity in structure and dynamics of SC trivalents was found for the first time. We assume that stretched centromeres were a marker of altered nuclear architecture in heterozygotes due to differences in the ancestral chromosomal territories of the parental species. Restructuring of the intranuclear organization and meiotic disturbances can contribute to the sterility of interspecific hybrids, and lead to the reproductive isolation of studied species.

We determined the body size and the size of molars (m/1) in mole voles (Ellobius talpinus Pallas 1770) collected in a laboratory colony (n = 35), in natural habitats (Orenburg oblast, n = 34; Chelyabinsk oblast, n = 42), and in two Holocene-dated horizons in Alekseevskaya Cave (n = 51; n = 66). It was found that the individual absolute age determined based on the molar root length (measured between the tooth neck and the end of the root) corresponds to the age intervals determined in labeled animals from natural habitats. The individual age of samples collected in Alekseevskaya Cave varied from several months to four years; the age group of one year was dominant, and animals younger than one year composed the second most abundant age group. The samples of the laboratory colony had the most pronounced morphological traits (greater height and length of the molar crown). We observed no significant differences in all the parameters studied between the animals collected in natural habitats and in the cave. Although there are no differences in the size parameters of the teeth, we cannot definitively conclude that the body size of mole voles in the Holocene was the same as that in recently collected samples, since the correlation of the body length and m/1 in animals from natural habitats and the laboratory colony was low.

In this study, we examined a total of 43 samples belonging to three Ellobius lutescens populations from Turkey, Iran and Nakhchivan, which are geographically separated by the Zagros, Tendürek and Alborz mountain ranges. We applied geometric morphometric methods (GMMs) to explore the differences in size and shape of the cranium and mandible. Indeed, we intriguingly found that the populations differed in cranium but not mandible size. Comparison of the Iranian and Turkish populations alone revealed morphological differences in the shape of the cranium and mandible that could be used as a barometer to predict the origin of individual animals. Importantly, our findings indicate that the Zagros and Tendürek mountain ranges may have acted as a barrier between these two populations, resulting in evolutionary divergence in these anatomical features. Consequently, we propose that within E. lutescens, subspecies including E. lutescens woosnami exists and in time, genetic, besides geographical barriers, may prevent subspecies from interbreeding with each other.

The weight parameters of the digestive tract, which characterize the size and filling of the fermentation chambers (forestomach and cecum), the mass of the walls and of the content of these organs, and the concentration and size composition of the fibers in the chyme have been determined in 12 individuals of the mole vole Ellobius talpinus born in captivity. The uneven development of the organs of the digestive tract of the mole vole is associated with the specificity of food specialization and does not serve as a compensatory mechanism. Large-size fractions of fiber masses dominate in the content of all organs; the specific content of the fibers is much lower in comparison with M. arvalis sp., which reflects the small impact of cellulose in the feeding of the mole vole. The bilocular discoglandular stomach of the mole vole, which is also typical for herbivorous rodents, is adapted to assimilate easy digestible fodder.

A typical feature of the populations of small mammals in different areas of the Saratov Trans-Volga region is the long-term and constantly increasing abundance of Sylvaemus uralensis and its wide distribution in a variety of habitats. The maximal abundance of S. uralensis is 72.0 individuals per 100 trap-days, and its share in catches is 73.0%. The abundance of Microtus arvalis tends to increase. It was shown that Lagurus lagurus has been found again in the steppe zone, on fallow lands and fields, after a prolonged depression in 2012–2013, the average density of species being 4.0 individuals per 100 trap-days. A high abundance of other typical steppe inhabitants (Allocricetulus eversmanni and Ellobius talpinus) was noted as well. The highest indices of species diversity (H = 1.18) are typical of fallow lands.

Morphometric parameters of hepatocytes have been studied in five small mammal species with different ecological specificity from two taxonomic groups: a surface-dwelling insectivore (the common shrew, Sorex araneus), a subterranean insectivore (the European mole, Talpa europaea), surface-dwelling rodents (the bank vole, Myodes glareolus and herb field mouse, Apodemus uralensis) and a subterranean rodent (the northern mole vole, Ellobius talpinus). The results show that the hepatocytes of the European mole differ markedly from those of all other species, being characterized by smaller size and nuclear area, low anisocytosis and anisokaryosis, the almost complete absence of binuclear cells and, on the other hand, high nuclear–cytoplasmic ratio and packing density. The common shrew differs from the European mole containing a relatively high proportion of binuclear hepatocytes, and the size of hepatocytes in this species is smaller than in rodents. Differences in the parameters of these cells between rodents are minimal or absent. The differences observed in the characteristics of hepatocytes between the above species cannot be explained solely by either taxonomic or ecological specificity. Comparative research involving a greater number of closely related species (at the levels of families and orders) is needed to reveal and evaluate the taxonomic and ecological specificity of hepatocytes in greater detail.

The involvement of chromosome changes in the initial steps of speciation is controversial. Here we examine diversification trends within the mole voles Ellobius, a group of subterranean rodents. The first description of their chromosome variability was published almost 40 years ago. Studying the G-band structure of chromosomes in numerous individuals revealed subsequent homologous, step-by-step, Robertsonian translocations, which changed diploid numbers from 54 to 30. Here we used a molecular cytogenetic strategy which demonstrates that chromosomal translocations are not always homologous, consequently, karyotypes with the same diploid number can carry different combinations of metacentrics. We further showed that at least three chromosomal forms with 2n = 34 and distinct metacentrics inhabit the Pamir-Alay mountains. Each of these forms independently hybridized with E. tancrei, 2n = 54, forming separate hybrid zones. The chromosomal variations correlate slightly with geographic barriers. Additionally, we confirmed that the emergence of partial or monobrachial homology appeared to be a strong barrier for hybridization in nature, in contradistinction to experiments which we reported earlier. We discuss the possibility of whole arm reciprocal translocations for mole voles. Our findings suggest that chromosomal translocations lead to diversification and speciation.

Restricted mobility and spatial isolation of social units in gregarious subterranean mammals ensure good defence mechanisms against parasites, which in turn allows for a reduction of immunity components. In contrast, a parasite invasion may cause an increased adaptive immune response. Therefore, it can be expected that spatial and temporal distribution of parasites within a population will correlate with the local variability in the host's immunocompetence. To test this hypothesis, the intra-population variability of a whipworm infestation and the humoral immune response to non-replicated antigens in mole voles (Ellobius talpinus Pall.), social subterranean rodents, was estimated. Whipworm prevalence in mole voles increased from spring to autumn, and this tendency was more pronounced in settlements living in natural meadows compared to settlements in man-made meadows. However, humoral immune response was lowest in animals from natural meadows trapped in autumn. Since whipworm infestation does not directly affect the immunity of mole voles, the reciprocal tendencies in seasonal dynamics and spatial distribution of whipworm abundance and host immunocompetence may be explained by local deterioration of habitat conditions, which increases the probability of an infestation.

The social environment can be stressful for at least some group members, resulting in elevated levels of glucocorticoid stress hormones (GC). Patterns of the relationships between social rank and GC levels vary between species. In carnivores, primates and birds that live in permanent cooperative groups, helpers do not usually display physiological indicators of stress. Very little is known about status-related GC differences within cooperative groups of rodents. In this laboratory study, we compared GC concentrations in dominant (fathers) and subordinate (natal sons) males of a cooperative subterranean vole, Ellobius tancrei. The assessment of adrenocortical activity by measuring urine glucocorticoid metabolites (UGM) was previously validated for this species through an ACTH challenge test. We observed clear peaks of UGM in the second or third urine samples taken after the administration of ACTH (lag time equal to 2.5–3 h). Thus, UGM is suitable to estimate physiological stress in Ellobius. Postpubertal sons living in natal groups had significantly higher UGM concentrations than their fathers. The average UGM levels of sons were positively associated with their ages and paternal body masses, and negatively associated with paternal ages. Hence, son-father interactions rather than just younger ages of sons appear to contribute to GC differences. The revealed pattern was not consistent with that reported for most cooperative species from other taxa, highlighting the importance of comparative studies.

The population of small mammals in different areas of the Saratov Trans-Volga region features long-term and constantly increasing numbers of Sylvaemus uralensis, and its wide distribution in a variety of habitats. The maximum density of S. uralensis is 72.0 ind. per 100 trap-days and the fraction in catches is 73.0%. A tendency is observed to an increasing numbers of Microtus arvalis. It is shown that after a prolonged depression in 2012-2013, Lagurus lagurus appeared in the steppe, on fallow lands and fields, the average density of species being 4.0 ind. per 100 trap-days. A high abundance of other typical steppe dwellers (Allocricetulus eversmanni and Ellobius talpinus) is noted as well. The highest indices of specific diversity (H = 1.18) are characteristic of fallow land.

The X and Y sex chromosomes of placental mammals show hallmarks of a tumultuous evolutionary past. The X Chromosome has a rich and conserved gene content, while the Y Chromosome has lost most of its genes. In the Transcaucasian mole vole Ellobius lutescens, the Y Chromosome including Sry has been lost, and both females and males have a 17,X diploid karyotype. Similarly, the closely related Ellobius talpinus, has a 54,XX karyotype in both females and males. Here, we report the sequencing and assembly of the E. lutescens and E. talpinus genomes. The results indicate that the loss of the Y Chromosome in E. lutescens and E. talpinus occurred in two independent events. Four functional homologs of mouse Y-Chromosomal genes were detected in both female and male E. lutescens, of which three were also detected in the E. talpinus genome. One of these is Eif2s3y, known as the only Y-derived gene that is crucial for successful male meiosis. Female and male E. lutescens can carry one and the same X Chromosome with a largely conserved gene content, including all genes known to function in X Chromosome inactivation. The availability of the genomes of these mole vole species provides unique models to study the dynamics of sex chromosome evolution.

Synaptonemal complex (SC) chains were revealed in semisterile intraspecific F1 hybrids of Ellobius tancrei Blasius, 1884 (2n = 49, NF=56 and 2n=50, NF=56), heterozygous for Robertsonian (Rb) translocations. Chains were formed by Rb submetacentrics with monobrachial homology. Chromosome synapsis in spermatocytes of these hybrids was disturbed, apparently because of the problematic release of the chromosomes from the SC chains. These hybrids suffer from low fertility, and our data support the opinion that this is because a formation of Rb metacentrics with monobrachial homology within different races of the same species might be an initial event for the divergence of chromosomal forms.

We present a first look at the microvertebrate fauna of the Middle Pleistocene site of Hummal in Central Syria. Some 2000 microvertebrate remains (1200 mammalian; 230 reptilian; 600 unidentified) were found in unit G/layer 17 by screen-washing sediments in an area of 4 m2. The following taxa have been identified: Reptilia: Agaminae indet., Gekkota indet., Lacertidae indet. (2–3 taxa), Eryx sp., Natricinae indet.; and Mammalia: Crocidurinae indet., Chiroptera indet., Lepus sp., Arvicolinae indet., Ellobius sp., Microtus sp., Murinae indet. (large form), Mus sp., Meriones sp., Gerbillus sp. The presence of Ellobius indicates a Middle Pleistocene age of the fauna. This genus does not occur in Syria today, but is recorded in Israel in Late Acheulean to Early Mousterian sites. In North Africa, the occurrence of this genus is restricted to the early part of the Middle Pleistocene. The ecological requirements of the nearest living relatives of the recorded taxa indicate mainly open habitats, but also the presence of vegetation and wet conditions, at least close to the site. Lepus, Ellobius, Meriones, Gerbillus, and Eryx live to various extents in steppes, semi-deserts, and deserts. Various extant species of large murids that come into consideration occupy a variety of wooded habitats, grassland and savannah and require the presence of water. Hummal offers the most detailed Pleistocene portrait yet of hominin palaeoenvironment in an area that, at the end of the Pleistocene, became known as part of the Fertile Crescent.

Dinopsyllus ellobius is considered a common and widespread flea in southern Africa and can act as a vector for plague. Due to differences in the interpretation of geographical variation in male sternite VIII, the taxonomy of the species is characterized by uncertainty. In an attempt to provide a better understanding of the systematics of D. ellobius, and also to provide new insights into the mechanisms that play a role in the diversification of the taxon, we sampled 830 small mammals at 31 localities throughout the distribution range of the parasite. Mitochondrial cytochrome c oxidase subunit II (COII) sequence data were generated for 151 D. ellobius specimens from 19 positive localities and this data set was supplemented with partial data derived from an elongation factor 1 alpha (EF1-α) intron. A parsimony haplotype network of the mtDNA and a Splitstree analyses revealed the existence of two genetic clades that were separated by 5.53% (± 1.78) sequence divergence. Although the nuclear DNA data were unresolved, significant size differences were detected between head, coxa, femur and tibia lengths of male individuals belonging to the two mtDNA lineages. The exact mechanisms that could have caused the diversification among lineages are not clear but the two lineages seem to be geographically separated and may have different ecological requirements. The present study strongly supports the notion that the two lineages are representative of D. ellobius (probably more associated with the host body and better adapted to mesic conditions) and Dinopsyllus abaris (probably more associated with the host nest and diverse climatic conditions) as originally proposed based on the single morphological character confined to male sternite VIII.

Ellobius talpinus was supposed to become extinct from the westernbank of the river Dnipro. Aft er 50 years the species was found again in Dnipropetrovsk and Kherson Region. The brief description of the current distribution of the northern mole vole to the west of the Dnipro is given. Altogether 11 localities were found. The rediscovered populations may be treated as endangered in the region.

Amdoparvoviruses (familyParvoviridae:genusAmdoparvovirus) infect carnivores, and are a major cause of morbidity and mortality in farmed animals. In this study, we systematically screened animal genomes to identify PVe disclosing a high degree of similarity to amdoparvoviruses, and investigated their genomic, phylogenetic and protein structural features. We report the first examples of full-length, amdoparvovirus-derived PVe in the genome of the Transcaucasian mole vole (Ellobius lutescens). Furthermore, we identify four further PVe in mammal and reptile genomes that are intermediate between amdoparvoviruses and their sister genus (Protoparvovirus) in terms of their phylogenetic placement and genomic features. In particular, we identify a genome-length PVe in the genome of a pit viper (Protobothrops mucrosquamatus) that is more like a protoparvovirus than an amdoparvovirus in terms of its phylogenetic placement and the structural features of its capsid protein (as revealed by homology modeling), yet exhibits characteristically amdoparvovirus-like genome features including: (i) a putative middle ORF gene; (ii) a capsid gene that lacks a phospholipase A2 (PLA2) domain; (iii) a genome structure consistent with an amdoparvovirus-like mechanism of capsid gene expression. Our findings indicate that amdoparvovirus host range has extended to rodents in the past, and that parvovirus lineages possessing a mixture of proto- and amdoparvovirus-like characteristics have circulated in the past. In addition, we show that PVe in the mole vole and pit viper encode intact, expressible replicase genes that have potentially been co-opted or exapted in these host species.

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