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Microtus (Sumeriomys) bifrons nov. sp. (Rodentia, Mammalia), a new vole in the French Upper Pleistocene identified at the Petits Guinards site (Creuzier-le-Vieux, Allier, France)

Microtus (Sumeriomys) bifrons nov. sp. (Rodentia, Mammalia), un nouveau campagnol pour le Pléistocène supérieur français, reconnu aux Petits Guinards (Creuzier-le-Vieux, Allier, France)
Marcel Jeannet and Laure Fontana
p. 59-77
This article is a translation of:
Microtus (Sumeriomys) bifrons nov. sp. (Rodentia, Mammalia), un nouveau campagnol pour le Pléistocène supérieur français, reconnu aux Petits Guinards (Creuzier-le-Vieux, Allier, France)

Abstracts

In a crept deposit, the only preserved part of the Magdalenian and Solutrean site of Les Petits Guinard (Allier, France), many thousands of Rodent bones and teeth have been identified. Among the numerous remains of local and boreal voles, one is unknown in France. It has been compared to the nearest morphological and geographical morphotypus: Microtus arvalis, agrestis, agrestis Leb. (temperate species), hyperboreus (boreal species), socialis and guentheri (eastern species). Biometrical data of these morphotypus (socialis excepted) have also been analysed. The whole data analysis clearly demonstrates that this vole of Les Petits Guinards is not one of the mentioned species, neither a sub-species. Never identified in France, living or fossilised, it is considered as a new species, called Microtus bifrons nov. sp.

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We are very grateful to Ms Margarita A. Erbajeva for her competent advice which enabled us to reduce our prospection field and save a lot of precious time. We also thank Christophe Petit for his help with the figures and the reviewers of this article for their helpful advice.

Introduction

1The Petits Guinards site (Creuzier-le-Vieux, Allier, France) was discovered in 1981 during reconstruction work on the road embankment bordering the right bank of the Allier (fig. 1) and was excavated in 2003 (Fontana et al. 2003a, 2003b). The geomorphological and archaeological study demonstrated the secondary position of the remains discovered inside a bulge deposit at the bottom of the slope. It attests to Palaeolithic occupations at the base of a limestone scarp, originally located at the top of a slope and dismantled at the beginning of the Holocene. The study also showed that the conserved part of the site was significantly displaced to 100 m lower down, and that the internal organization of the site stratigraphy did not undergo any major sedimentary reworking. Nonetheless, it was not possible to differentiate between any potential layers in the archaeological level, which reaches a thickness of 60 to 120 cm, due to the type of deposit (heterometric blocks in a silty matrix). The thirteen radiocarbon dates on bone and dental remains (food waste and objects in reindeer antler) place the occupations in a long time period ranging from 19 600 to 10 300 uncal BP. Several flints provide clear presence of a Solutrean occupation (Fontana et al. 2013), but most of the lithic industry is Magdalenian, as is the industry in hard animal matter (Fontana et al. 2003a, 2003b). The abundance of the latter (Fontana & Chauvière 2009; Chauvière et al. 2006) and the presence of the Solutrean are two original features of this Upper Palaeolithic site located in the Massif Central. The abundance and the excellent conservation of the bone remains are also remarkable. The 100 000 rodent remains include numerous temperate and boreal species, providing evidence of a continental climate: the mole (Talpa europaea), the common shrew (Sorex araneus), the greater white-toothed shrew (Crocidura russula), the field vole (Microtus agrestis) and emblematic species such as the Arctic lemming (Dicrostonyx torquatus), the Tundra vole (Microtus oeconomus), the Narrow-headed vole (Microtus gregalis), the Northern birch mouse (Sicista betulina) and the very abundant Russet ground squirrel (Spermophilus major) (fig. 2). Most of the rodent remains belong to six species of voles (including five Microtus), but some vole molars present differences, raising questions as to their affiliation to one of the six determined microtine species. The aim of this study is thus to identify the species represented by these dental remains.

Figure 1 - Les Petits Guinards, Creuzier- le-Vieux. Map of site location.

Figure 1 - Les Petits Guinards, Creuzier- le-Vieux. Map of site location.

Figure 2 - Microvertebrate species from Les Petits Guinards.

Figure 2 - Microvertebrate species from Les Petits Guinards.

1 – Dental morphology and morphometric analysis

1.1 – Dental morphology

2The dental remains of small voles from the Petits Guinards deposits represent a minimum number of 430 individuals. The terminology of the occlusal surface elements of the molars is presented in figure 3. Sixteen m/1 and one M3/ present significant differences, and are at the limitations of the initial characteristics of the Microtus genus1. The general aspect of these molars is similar to the Microtus genus, but several characteristics sporadically observed in diverse species are combined here on members of this group with remarkable constancy and intensity: in particular the acuity of the projecting angles, the hermetic closing of the triangles and their very marked bucco-jugal symmetry. But, above all, although eight m1 present five triangles and two present six (fig. 4), surprisingly, the six other m1 bear seven closed triangles (sometimes converging to form a deformed rhombus tending to split), which only occurs in exceptional cases in other microtines. In addition, among the eight m1 with five triangles, each group of five triangles is preceded by a rhombus deformed by the penetration of the internal angle a7, which tends to create two new closed triangles: these m1 are thus part of the group of molars with seven closed triangles. Forms 7 and 17 (cf. fig. 4), with a very open rhombus, are separate from the other types. Lastly, a tight constriction isolates the anterior loop, which is extended by a characteristic spur on the external surface (t8), which never occurs elsewhere (cf. fig. 4, n. 13, 14).

3What species do these M1 belong to? They could be attributed to two of the five microtine species identified at Petits Guinards (cf. fig. 2); Microtus arvalis and Microtus agrestis, which are morphologically similar to this new arrival. M. arvalis differs by its smaller size, less acute bulging angles, practically no dissymmetry, a less developed and more rounded t8 (fig. 5). As for M. agrestis, the relatively frequent presence of six closed and angular triangles is similar to indeterminate Microtus (cf. fig. 5c), as it never presents seven closed triangles or such a bulging t8. Figure 6 presents the anterior loop of present-day voles from Finistère (where Microtus arvalis is naturally absent) and provides evidence of this; it associates a wide variety of chosen forms, from the most simple to the most complex, in order to avoid all confusion with the common vole (M. arvalis), its closest relative, naturally absent from Brittany.

4This indeterminate vole differs from local microtines, but may perhaps be compared to boreal species issued from migratory populations, as such species are present in Petits Guinards? The comparison with Microtus hyperboreus Vinogradov 1933, the North Siberian vole, issued from the Lena Delta with a territory extending from the north of the Ural to the Jamal Peninsula appears to be an obvious point of comparison as it is the only Boreal microtine to be naturally associated with Dicrostonyx torquatus. Its m/1 bears five to six closed angular and dissymmetrical triangles (occasionally seven), which relate it to indeterminate Microtus. However, the t8 has no jugal spur and no constriction of the anterior loop is visible. In addition, the M. hyperboreus M3 bears an additional internal protruding angle. These differences appear to be sufficiently significant to rule out an attribution of the indeterminate microtine from Petits Guinards to this subarctic species.

5It thus seems impossible to consider the indeterminate microtine from Petits Guinards as a subspecies of one of the presented microtines as the morphological differences are too marked and are beyond the variability of the Microtus genus. If we did not take account of these differences, we would have to redefine the whole systematics of the genus. The best example of this is the similarity between M. arvalis and M. agrestis, leading many authors to regroup them under the double label M. arvalis-agrestis (Chaline 1972; Desclaux et Defleur 1997; Sese 2005; Cuenca-Bescos et al. 2010). Yet, the criteria presented here for describing this new Microtus are much clearer than those likely to separate the two afroementioned species. This observation is all the more pertinent for the other analysed microtines. Do the morphometric data confirm this conclusion?

Figure 3 - Ondatra zibethicus.Terminology example of the occlusal surface of vole molars (from Hibbard, 1950, fig.16). a/ m1-m3 G; b/ M1-M3 G. AC: anterior cap; ACC: anteroconid complex; AL: anterior loop; BRA: buccal re- entrant angle; BSA: buccal salient angle; LRA: lingual re- entrant angle; LSA: lingual salient; PC: Posterior cap; PL: Posterior loop. TTC: trigonid-talonid complex.

Figure 3 - Ondatra zibethicus.Terminology example of the occlusal surface of vole molars (from Hibbard, 1950, fig.16). a/ m1-m3 G; b/ M1-M3 G. AC: anterior cap; ACC: anteroconid complex; AL: anterior loop; BRA: buccal re- entrant angle; BSA: buccal salient angle; LRA: lingual re- entrant angle; LSA: lingual salient; PC: Posterior cap; PL: Posterior loop. TTC: trigonid-talonid complex.

Figure 4 - Microtus bifrons - Les Petits Guinards, Creuzier-le-Vieux. Scale: 1 mm.

Figure 4 - Microtus bifrons - Les Petits Guinards, Creuzier-le-Vieux. Scale: 1 mm.

Figure 5 - a/ Microtus arvalis; m1, m2, m3 G; b/ Microtus arvalis: M1, M2, M3 G. ; c/ Microtus agrestis: m1, m2, m3 G; d/ Microtus agrestis: M1, M2, M3 G. Grotte des Romains, (Virignin, Ain) – Magdalenian (Excavations by R. Desbrosse).

Figure 5 - a/ Microtus arvalis; m1, m2, m3 G; b/ Microtus arvalis: M1, M2, M3 G. ; c/ Microtus agrestis: m1, m2, m3 G; d/ Microtus agrestis: M1, M2, M3 G. Grotte des Romains, (Virignin, Ain) – Magdalenian (Excavations by R. Desbrosse).

Figure 6 – Present-day Microtus agrestis from Finistère. Anterior complex from m1. Varied types are classified from 1 to 14 following the evolution of the closing of t6 and progressive development of t8 on the anterior loop (N.B.: Microtus arvalis is currently absent from Finistère, Côtes-d’Armor and Morbihan).

Figure 6 – Present-day Microtus agrestis from Finistère. Anterior complex from m1. Varied types are classified from 1 to 14 following the evolution of the closing of t6 and progressive development of t8 on the anterior loop (N.B.: Microtus arvalis is currently absent from Finistère, Côtes-d’Armor and Morbihan).

Figure 7 – Position of biometric points taken on Microtus M/1 (adapted from Van der Meulen 1973).

Figure 7 – Position of biometric points taken on Microtus M/1 (adapted from Van der Meulen 1973).

Figure 8 - Comparison of morphometric values for the unidentified microtine from Petits Guinards and four other Microtines.

Figure 8 - Comparison of morphometric values for the unidentified microtine from Petits Guinards and four other Microtines.

Figure 9 - Comparison of Microtus biometric data from Les Petits Guinards and four other microtines (M. arvalis, M. agrestis, M. agrestis Leb., M. hyperboreus), using the Mann-Whitney test. In grey: non-significant differences (p >0.01)

Figure 9 - Comparison of Microtus biometric data from Les Petits Guinards and four other microtines (M. arvalis, M. agrestis, M. agrestis Leb., M. hyperboreus), using the Mann-Whitney test. In grey: non-significant differences (p >0.01)

1.2 - Morphometry

6The m/1 metric data from the five analysed morphotypes (from Petits Guinards: M. arvalis, M. agrestis and M. indet.; from the current collections: M. hyperboreus and M. agrestis Leb.) are grouped together in the annexed tables (annexes 1 to 5). We analysed seven variables for these five morphotypes (fig. 7): the length of the M/1, the ratio between the constriction of the anterior loop (b2) and the internal width of its base (W2) (following the Van der Meulen standards, 1973), the reduction ratio (b2/c2) for the two isthmuses of the anterior loop, the c2/W2 ratio, the asymmetry of the anterior loop (asym), the asymmetry/anterior complex ratio (ACC), the asymmetry of the trigonid (Wn/Wr). As the aim of this is to compare the indeterminate Microtus from Petits Guinards (called MPG) to the other described microtines, we tested the differences between Microtus indet. and each morphotype for each variable, using the Mann-Withney test. The results are presented in figures 8 and 9.

7The length of the M/1 from MPG is significantly greater than that of three of the four microtines, with the exception of M. agrestis, like for the asymmetry/anterior complex ratio (ACC). The reduction ratio (b2/c2) is also significantly different between MPG and three microtines, with the exception of M. hyperboreus, like for the c2/W2 ratio. For c2/W2, the closing of t6 depends on the c2 value and can sometimes be considered as an evolutionary character of the molar which tends to get longer through the acquisition of an additional closed triangle. This is naturally the case for MPG which generally has seven closed triangles, as shown in the figure. The figure also shows the discrepancy of the c2/w2 index towards the lowest MPG values. According to the Mann-Withney test (cf. fig. 9), it is distinct from the two M. agrestis and M. arvalis types. The ratio between the constriction of the anterior loop and the internal width of the base (b2/W2) is significantly different between MPG and the four microtines. As for the asymmetry of the trigonid (Wn/Wr) and the asymmetry of the anterior loop (Asym), only the difference between MPG and M. agrestis Leb. is significant. In spite of a visual dissymmetric appearance of the anterior loop, the recorded values are uniformly incorporated in the mass of the other taxa, which is rather unexpected. We can consider this as confirmation of the attribution of MPG to the Microtus genus. The asymmetry of the (Wn/Wr) trigonid is only evident for the Breton Microtus agrestis (C). The visual aspect of this parameter thus appears to be illusory in a multi-specific association.

8All of these differences illustrate three main facts. First of all, the microtine from Petits Guinards can be statistically differentiated on the basis of one parameter (the b2/W2 ratio). For four of the six other variables (length, b2/c2, c2/w2, ACC), the difference between MPG and the four other microtines is significant for three of them, which signifies that for each of these four variables, MPG is only similar to one of the four microtines, that is M. hyperboreus in two cases and M. agrestis in two cases. The two other variables (the two asymmetries) are only significantly different in one case, which is M. agrestis Leb. Furthermore, a single morphotype systematically presents significant differences with MPG (for the seven variables, cf. fig. 9); that is M. agrestis Leb.

9Consequently, none of the four morphotypes is significantly similar to MPG, according to the measured parameters, which confirms that this microtine is a different species to the local microtine species and the tested boreal species. The two dominant criteria are the presence of seven closed triangles on the M/1 (rarely six or eight), and the large size. The constriction of the anterior loop and the spurred t8 are also characteristic. On the other hand, the visually evident asymmetry does not resist to the biometric test. What species do these M/1 thus belong to?

2 – A Near Eastern vole in the Upper French Pleistocene?

2.1 - Microtus socialis and Microtus guentheri

10The indeterminate Microtus from Petits Guinards does not thus appear to be a local type vole or a Siberian migrant, which incites us to seek its origin in lower latitudes and more rigorous climates. The latitudinal and climatic convergence could be compared to the gopher, which is very abundant at Petit Guinards and which extends to the semi-arid and steppe-like regions of south-western Asia. At this stage, only the literature provides information on this, without taking regional faunal associations into consideration. S. I. Ognev (1950) regrouped all the sub-generic mid-oriental forms of Chilotus Baird 1857 (North American microtine sub-genus) under the single specific name of Microtus socialis, the social vole, thereby demoting them to sub-specific status. According to that author, the domain of M. socialis extends from the north of the Black Sea to inland Mongolia and it also lives in low altitude semi-desert zones (fig. 10), like the gopher. The application of mitochondrial research (Yigit & Çolak 2002; Golenischchev et al. 2002a, 2000b; Jaarola et al. 2004; Yigit et al. 2012) modified this model by placing the the so-called subspecies into two main groups of species2: Microtus guentheri and M. socialis. These works also revised the distribution of M. socialis (by S.I. Ognev 1950), which extended over the geographic zone in the north Caucasus region and a large strip of the Black Sea to Kazakhstan (Corbet 1978; Gromov & Erbajeva 1995; Yigit et al. 2003; Yigit et al. 2012) (cf. fig. 10). This species was sporadically recorded in several Near Eastern sites (Kowalski 1958). As for M. guentheri, it extends to the Black Sea in the south, to Kirghizistan in the east (Lac Ala Kul), to the south (Lebanon, Syria, Israel and isolated occurrences in Cyrenaica) and to the west (Thrace and Bulgaria). For some researchers, its domain extends to the Balkans.

11The two species differ by their dental morphology and size.

12Most of the m/1 from these two species are made up of five closed triangles, rarely six, but those of Microtus socialis3 are the only ones with seven closed triangles and sometimes six; they are also the only ones with a protruding t8 (Ognev 1950, fig. 11). In addition, we observe the frequent presence on the posterior end of a protuberance that can attain the shape of a loop on the M/2, which is well established for M. agrestis (cf. fig. 5d). On the M/1, this shape is never so developed and is limited to a more or less voluminous angle (cf. fig. 11). The M/3 displays four bulging lingual and three external angles, but these criteria do not seem to be retained for determinations, probably on account of their instability.

13M. guentheri (Danford & Alston, 1880) (fig. 12, 13, 14) differs from M. socialis by a larger molar size (M. socialis is the smallest species of the Sumeriomys sub-genus), orange-coloured incisors (Yigit & Çolak 2002) and the fact that the two upper molars do not have posterior agrestoid formations (Baydemir & Duman 2009).

14M. guentheri can be separated from M. socialis on the basis of its large size. G. Storch (1988) indicates, for the M/ of M. guentheri, an average length of 330 (300-360), which is higher than that of MPG (306; 280-345), but no M/1 dimensions figure in the published studies.

Figure 10 - Distribution map of Microtus socialis and Microtus guentheri.

Figure 10 - Distribution map of Microtus socialis and Microtus guentheri.

2.2 - The microtine from Petits Guinards: Microtus bifrons nov. sp.

15It thus seems that the dental morphology of the microtine from Petits Guinards assimilates it to Microtus socialis, as described by Ognev (1950) if we exclude the adventive formations extending the posterior end of the first two upper molars. These protuberances, with instable size and presence, frequently characterize this taxon. For this point, we follow the denomination defined by Ognev for this very characteristic morphology, which is not found anywhere else. The seven closed reference triangles are present on the microtine from Petits Guinards and M. socialis, but the latter is too small for MPG to be a sub-species. The larger size and morphology of MPG relate it to the sub-genus Sumeriomys (fig. 15), but differentiate it from the two morphotypes; one on account of its shape (M. guentheri), and the other its size (M. socialis), although it conserves a double facies. It is thus difficult to name the vole from Petits Guinards.

16The excessive size could be an evolutionary sign of the species in a more favourable environment than the semi-desert type conditions of its initial biotope. The “socialis” form described by Ognev (1950) changed little but its size increased considerably to reach the size of M. guentheri (while losing the agrestic formations of the first two upper molars). In this way, this new vole bears the marks of two major Mid-Eastern Sumeriomys taxa. This is why we employ the adjective “bifrons”, denoting the double facies of the first lower molar. The characteristics are the following:

17MICROTUS BIFRONS nov. sp.
FAMILY: Cricetides
SUB-FAMILY: Microtines
GENUS: Microtus SCHRANK, 1798
SUB-GENUS: Sumeriomys ARGYROPULO, 1933
HOLOTYPE: m1G n° CVPG 57-1248a (fig. 16)
LOCALITY-TYPE: Creuzier-le-Vieux (Allier); locality Les Petits Guinards
TYPE-LEVEL : The whole of the Upper Pleniglacial with Solutrean and Magdalenian industries.
PROBABLE AGE: 20 000 – 11 000 uncal BP
ORIGIN OF THE NAME: from the Latin bifrons: Latin term signifying double face; used on account of the double facies of the m1: Microtus socialis for its polymorphic dental pattern (six to seven closed triangles, rarely five), and Microtus guentheri for its large size and the absence of agrestis type upper molars.
DIAGNOSIS: Large-sized vole with six to seven highly dissymmetrical and acute angled closed triangles on the m1. Highly dissymmetrical anterior loop and voluminous spur-shaped t8.
PARATYPES: M3D with 3 closed triangles, five external protruding angles and four internal protruding angles (cf. fig. 16).

18We now have better knowledge of the current distribution of M. socialis and guentheri (see above), but can data relating to their former geographic distribution contribute to our understanding of the presence of a vole such as M. bifrons in the Massif Central between 20 000 and 11 000 uncal BP (MIS 2 and the Tardiglacial) and probably derived from Eastern species? Unfortunately, such mentions are rare and are dispersed in space and time (Kowalski 1958; Tchernov 1968; Storch 1988; Gromov & Erbajeva 1995; Helmer et al. 1998; Santel & von Koenigswald 1998; Khenzykhenova et al. 2011; Markova 2011; Maul et al. 2011; Popova 2004). These two Microtus are indicated from the Middle Pleistocene onwards in sites in the Near East, Anatolia, in the north of the Caucasus and in Thrace. The hypothesis of the migration of one or two of these species from Turkey by successive waves since the Middle Pleistocene during the drying of the Bosporus Strait is thus currently preferred.

Figure 11 - Microtus socialis after S.I. Ognev, 1950 - a/ 1- 3: m1D; 1 – Northern Caucasus; M. s. parvus; 2/ Ala Kul Lake (Khirgyzistan): M. s. gravesi; 3/ Kaine-Kassyr (Turkmenistan, near the Iranian border: M. s. paradoxus (Ognev 1950, fig. 163). b/ Structure variations in Microtus socialis molars. 1-3: Salk region; 4: Bakou region; 5-6 Kopet, region Dag. 1,3,4,5,6 : M1-M3D; 2 : m1-m3D. (Ognev 1950, fig. 162). No scale.

Figure 11 - Microtus socialis after S.I. Ognev, 1950 - a/ 1- 3: m1D; 1 – Northern Caucasus; M. s. parvus; 2/ Ala Kul Lake (Khirgyzistan): M. s. gravesi; 3/ Kaine-Kassyr (Turkmenistan, near the Iranian border: M. s. paradoxus (Ognev 1950, fig. 163). b/ Structure variations in Microtus socialis molars. 1-3: Salk region; 4: Bakou region; 5-6 Kopet, region Dag. 1,3,4,5,6 : M1-M3D; 2 : m1-m3D. (Ognev 1950, fig. 162). No scale.

Figure 12 - Microtus guentheri. a/ 1-11: Lower cheek teeth. Acheulean. Oum Qatafa Cave (Israel). Tchernov 1968. b/ 1-6: Upper cheek teeth. Acheulean. Oum Qatafa Cave (Israel). Tchernov 1968. c/ 1: Upper cheek teeth (M1-M3D). Upper Levallloiso-Mousterian. Kebara Cave (Israel). c1-5: Upper cheek teeth G. recent (Tchernov 1968). No scale.

Figure 12 - Microtus guentheri. a/ 1-11: Lower cheek teeth. Acheulean. Oum Qatafa Cave (Israel). Tchernov 1968. b/ 1-6: Upper cheek teeth. Acheulean. Oum Qatafa Cave (Israel). Tchernov 1968. c/ 1: Upper cheek teeth (M1-M3D). Upper Levallloiso-Mousterian. Kebara Cave (Israel). c1-5: Upper cheek teeth G. recent (Tchernov 1968). No scale.

Figure 13 - Microtus guentheri. a & b/ M3D; c & d/ m1-m3D. Kirikkale Province (Turkey). Baydemir & Duman (2009). No scale.

Figure 13 - Microtus guentheri. a & b/ M3D; c & d/ m1-m3D. Kirikkale Province (Turkey). Baydemir & Duman (2009). No scale.

Figure 14 - Microtus guentheri. 10-12: m1G; 13-15: M3G. Microtus arvalis. 13-15: m1; 19-20: M3D. Karain B. (Turkey). Storch (1988). No scale.

Figure 14 - Microtus guentheri. 10-12: m1G; 13-15: M3G. Microtus arvalis. 13-15: m1; 19-20: M3D. Karain B. (Turkey). Storch (1988). No scale.

Figure 15 - Sub-genus Sumeriomys – m1D (A, C, E, G, J, K, M, O, R, T, V, X, et Z) & m3D (B, D, H, I, L, N, P, S, U,W,Y, Z& ZZ). A & B: Microtus socialis socialis (Gur’ev, Kazakhstan); C& D: M. s. nikolajevi (Kuyuk-Tuk Island, Ukraine); E & F: M. s. binominatus (Tbilissi, Georgia); G & H: M. s. goriensis (Tamarasheni, Georgia); I & J: M. s. goriensis (Tamarasheni, Géorgia); K & L: M. schidlovvkii (Nalband, Armenia); M & N: M. parvus (Divnoye-Stavropol, Russia); O & P: M. paradoxus (Ashkhabad, Turkmenistan); T & U: M. s. gravesi (Betpakdala Steppe, Kazakhstan); V & W: M. guentheri stranjensis (Sozopol, Bulgaria); X & Y: M. s. zaitsevi (Holotype – Bakou, Azerbaijan); Z & ZZ: M. s. aristovi (Holotype – Veysalli, Azerbaijan). Golenischchev et al. 2002a. Scale 1mm.

Figure 15 - Sub-genus Sumeriomys – m1D (A, C, E, G, J, K, M, O, R, T, V, X, et Z) & m3D (B, D, H, I, L, N, P, S, U,W,Y, Z& ZZ). A & B: Microtus socialis socialis (Gur’ev, Kazakhstan); C& D: M. s. nikolajevi (Kuyuk-Tuk Island, Ukraine); E & F: M. s. binominatus (Tbilissi, Georgia); G & H: M. s. goriensis (Tamarasheni, Georgia); I & J: M. s. goriensis (Tamarasheni, Géorgia); K & L: M. schidlovvkii (Nalband, Armenia); M & N: M. parvus (Divnoye-Stavropol, Russia); O & P: M. paradoxus (Ashkhabad, Turkmenistan); T & U: M. s. gravesi (Betpakdala Steppe, Kazakhstan); V & W: M. guentheri stranjensis (Sozopol, Bulgaria); X & Y: M. s. zaitsevi (Holotype – Bakou, Azerbaijan); Z & ZZ: M. s. aristovi (Holotype – Veysalli, Azerbaijan). Golenischchev et al. 2002a. Scale 1mm.

Figure 16 - Microtus bifrons (Petits Guinards). a/Holotype: m1 left. b/ Paratype: m1 right. c/ Paratype: M3 right. Scale 1 mm.

Figure 16 - Microtus bifrons (Petits Guinards). a/Holotype: m1 left. b/ Paratype: m1 right. c/ Paratype: M3 right. Scale 1 mm.

Conclusion

19Given the major differences between this new form from Petits Guinards and the described microtine species, it seems logical to attribute a specific status to the vole from Petits Guinards in keeping with the developed criteria. These differences were identified on the m1, the most obvious being the increased number of closed triangles and the marked development of the t8 resulting in increased size. The combined association of the dental morphology of Microtus socialis (Pallas, 1773) and the large size of Microtus guentheri (Danford et Alston, 1880) resulted in the name Microtus bifrons nov. sp., indistinctly associating the double facies of both taxa. Due to the focus on North Siberian continental species, we tend to forget that migrations can also occur from east to west, along the same latitudinal gradient, and the association of micromammals from Petits Guinards is an invaluable reminder of this. The history of this lineage of microtines and the arrival of Microtus bifrons in Europe remain to be determined, through the continued study of micromammal remains, which are at times very abundant and very well conserved in the sediments accumulated in caves or rock shelters.

Annex 1 – Metric data from the Microtus arvalis m1 from Petits Guinards (Massif Central).

Annex 1 – Metric data from the Microtus arvalis m1 from Petits Guinards (Massif Central).

Annex 2 – Metric data from Microtus agrestis m1 from Petits Guinards (Massif Central).

Annex 2 – Metric data from Microtus agrestis m1 from Petits Guinards (Massif Central).

Annex 3 – Metric data from the present-day Microtus agrestis m1 (Brittany).

Annex 3 – Metric data from the present-day Microtus agrestis m1 (Brittany).

Annex 4 – Metric data from the Microtus bifrons m1 from Petits Guinards (Massif Central).

Annex 4 – Metric data from the Microtus bifrons m1 from Petits Guinards (Massif Central).

Annex 5 – Metric data from present-day Microtus hyperboreus m1 (Yakutie).

Annex 5 – Metric data from present-day Microtus hyperboreus m1 (Yakutie).
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Bibliography

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List of illustrations

Title Figure 1 - Les Petits Guinards, Creuzier- le-Vieux. Map of site location.
URL http://journals.openedition.org/paleo/docannexe/image/3026/img-1.png
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Title Figure 2 - Microvertebrate species from Les Petits Guinards.
URL http://journals.openedition.org/paleo/docannexe/image/3026/img-2.png
File image/png, 39k
Title Figure 3 - Ondatra zibethicus.Terminology example of the occlusal surface of vole molars (from Hibbard, 1950, fig.16). a/ m1-m3 G; b/ M1-M3 G. AC: anterior cap; ACC: anteroconid complex; AL: anterior loop; BRA: buccal re- entrant angle; BSA: buccal salient angle; LRA: lingual re- entrant angle; LSA: lingual salient; PC: Posterior cap; PL: Posterior loop. TTC: trigonid-talonid complex.
URL http://journals.openedition.org/paleo/docannexe/image/3026/img-3.png
File image/png, 32k
Title Figure 4 - Microtus bifrons - Les Petits Guinards, Creuzier-le-Vieux. Scale: 1 mm.
URL http://journals.openedition.org/paleo/docannexe/image/3026/img-4.png
File image/png, 135k
Title Figure 5 - a/ Microtus arvalis; m1, m2, m3 G; b/ Microtus arvalis: M1, M2, M3 G. ; c/ Microtus agrestis: m1, m2, m3 G; d/ Microtus agrestis: M1, M2, M3 G. Grotte des Romains, (Virignin, Ain) – Magdalenian (Excavations by R. Desbrosse).
URL http://journals.openedition.org/paleo/docannexe/image/3026/img-5.png
File image/png, 24k
Title Figure 6 – Present-day Microtus agrestis from Finistère. Anterior complex from m1. Varied types are classified from 1 to 14 following the evolution of the closing of t6 and progressive development of t8 on the anterior loop (N.B.: Microtus arvalis is currently absent from Finistère, Côtes-d’Armor and Morbihan).
URL http://journals.openedition.org/paleo/docannexe/image/3026/img-6.png
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Title Figure 7 – Position of biometric points taken on Microtus M/1 (adapted from Van der Meulen 1973).
URL http://journals.openedition.org/paleo/docannexe/image/3026/img-7.png
File image/png, 58k
Title Figure 8 - Comparison of morphometric values for the unidentified microtine from Petits Guinards and four other Microtines.
URL http://journals.openedition.org/paleo/docannexe/image/3026/img-8.png
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Title Figure 9 - Comparison of Microtus biometric data from Les Petits Guinards and four other microtines (M. arvalis, M. agrestis, M. agrestis Leb., M. hyperboreus), using the Mann-Whitney test. In grey: non-significant differences (p >0.01)
URL http://journals.openedition.org/paleo/docannexe/image/3026/img-9.png
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Title Figure 10 - Distribution map of Microtus socialis and Microtus guentheri.
URL http://journals.openedition.org/paleo/docannexe/image/3026/img-10.png
File image/png, 264k
Title Figure 11 - Microtus socialis after S.I. Ognev, 1950 - a/ 1- 3: m1D; 1 – Northern Caucasus; M. s. parvus; 2/ Ala Kul Lake (Khirgyzistan): M. s. gravesi; 3/ Kaine-Kassyr (Turkmenistan, near the Iranian border: M. s. paradoxus (Ognev 1950, fig. 163). b/ Structure variations in Microtus socialis molars. 1-3: Salk region; 4: Bakou region; 5-6 Kopet, region Dag. 1,3,4,5,6 : M1-M3D; 2 : m1-m3D. (Ognev 1950, fig. 162). No scale.
URL http://journals.openedition.org/paleo/docannexe/image/3026/img-11.png
File image/png, 1.4M
Title Figure 12 - Microtus guentheri. a/ 1-11: Lower cheek teeth. Acheulean. Oum Qatafa Cave (Israel). Tchernov 1968. b/ 1-6: Upper cheek teeth. Acheulean. Oum Qatafa Cave (Israel). Tchernov 1968. c/ 1: Upper cheek teeth (M1-M3D). Upper Levallloiso-Mousterian. Kebara Cave (Israel). c1-5: Upper cheek teeth G. recent (Tchernov 1968). No scale.
URL http://journals.openedition.org/paleo/docannexe/image/3026/img-12.png
File image/png, 114k
Title Figure 13 - Microtus guentheri. a & b/ M3D; c & d/ m1-m3D. Kirikkale Province (Turkey). Baydemir & Duman (2009). No scale.
URL http://journals.openedition.org/paleo/docannexe/image/3026/img-13.png
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Title Figure 14 - Microtus guentheri. 10-12: m1G; 13-15: M3G. Microtus arvalis. 13-15: m1; 19-20: M3D. Karain B. (Turkey). Storch (1988). No scale.
URL http://journals.openedition.org/paleo/docannexe/image/3026/img-14.png
File image/png, 15k
Title Figure 15 - Sub-genus Sumeriomys – m1D (A, C, E, G, J, K, M, O, R, T, V, X, et Z) & m3D (B, D, H, I, L, N, P, S, U,W,Y, Z& ZZ). A & B: Microtus socialis socialis (Gur’ev, Kazakhstan); C& D: M. s. nikolajevi (Kuyuk-Tuk Island, Ukraine); E & F: M. s. binominatus (Tbilissi, Georgia); G & H: M. s. goriensis (Tamarasheni, Georgia); I & J: M. s. goriensis (Tamarasheni, Géorgia); K & L: M. schidlovvkii (Nalband, Armenia); M & N: M. parvus (Divnoye-Stavropol, Russia); O & P: M. paradoxus (Ashkhabad, Turkmenistan); T & U: M. s. gravesi (Betpakdala Steppe, Kazakhstan); V & W: M. guentheri stranjensis (Sozopol, Bulgaria); X & Y: M. s. zaitsevi (Holotype – Bakou, Azerbaijan); Z & ZZ: M. s. aristovi (Holotype – Veysalli, Azerbaijan). Golenischchev et al. 2002a. Scale 1mm.
URL http://journals.openedition.org/paleo/docannexe/image/3026/img-15.png
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Title Figure 16 - Microtus bifrons (Petits Guinards). a/Holotype: m1 left. b/ Paratype: m1 right. c/ Paratype: M3 right. Scale 1 mm.
URL http://journals.openedition.org/paleo/docannexe/image/3026/img-16.png
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Title Annex 1 – Metric data from the Microtus arvalis m1 from Petits Guinards (Massif Central).
URL http://journals.openedition.org/paleo/docannexe/image/3026/img-17.png
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Title Annex 2 – Metric data from Microtus agrestis m1 from Petits Guinards (Massif Central).
URL http://journals.openedition.org/paleo/docannexe/image/3026/img-18.png
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Title Annex 3 – Metric data from the present-day Microtus agrestis m1 (Brittany).
URL http://journals.openedition.org/paleo/docannexe/image/3026/img-19.png
File image/png, 56k
Title Annex 4 – Metric data from the Microtus bifrons m1 from Petits Guinards (Massif Central).
URL http://journals.openedition.org/paleo/docannexe/image/3026/img-20.png
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Title Annex 5 – Metric data from present-day Microtus hyperboreus m1 (Yakutie).
URL http://journals.openedition.org/paleo/docannexe/image/3026/img-21.png
File image/png, 6.5k
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References

Bibliographical reference

Marcel Jeannet and Laure Fontana, « Microtus (Sumeriomys) bifrons nov. sp. (Rodentia, Mammalia), a new vole in the French Upper Pleistocene identified at the Petits Guinards site (Creuzier-le-Vieux, Allier, France) », PALEO, 26 | 2015, 59-77.

Electronic reference

Marcel Jeannet and Laure Fontana, « Microtus (Sumeriomys) bifrons nov. sp. (Rodentia, Mammalia), a new vole in the French Upper Pleistocene identified at the Petits Guinards site (Creuzier-le-Vieux, Allier, France) », PALEO [Online], 26 | 2015, Online since 26 April 2016, connection on 09 December 2018. URL : http://journals.openedition.org/paleo/3026

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About the authors

Marcel Jeannet

UMR 7269 LAMPÉA, BP 647, Maison Méditerranéenne des Sciences de l’Homme, 5 rue du château de l’horloge - BP 647, FR‑13094 Aix-en-Provence cedex - m.jeannet.arpa.mf@wanadoo.fr

By this author

Laure Fontana

(b) CNRS, UMR 7401 ArScAn, Archéologies environnementales. Maison de l‘Archéologie et de l’Ethnologie R. Ginouvès, 21 Allée de l’Université, FR-92023 Nanterre cedex - laure.fontana@mae.u-paris10.fr

By this author

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