A new species of Blarinoides (Soricidae, Mammalia), Blarinoides aliciae, is described from the Pliocene of Tollo de Chiclana (Guadix Basin, southeastern Spain). It is notably smaller than the single species of this genus known until now, B. mariae. B. aliciae is recorded from several other Spanish localities, but the sample from Tollo de Chiclana-3 is the most numerous. B. aliciae proves the simultaneous existence of two different species of Blarinoides in Europe during the Pliocene: one of large size in central and eastern Europe, and another one of small size in Iberia.
Une nouvelle espèce de Blarinoides (Soricidae, Mammalia), Blarinoides aliciae, est décrite du Pliocène de Tollo de Chiclana (dépression de Guadix, Espagne sud-orientale). Elle est notablement plus petite que la seule espèce du genre connue jusqu’à maintenant, B. mariae. B. aliciae est documentée dans plusieurs autres gisements espagnols, mais l’échantillon de Tollo de Chiclana-3 est le plus abondant. B. aliciae démontre l’existence simultanée de deux espèces de Blarinoides en Europe au Pliocène : une première, de grande taille, en Europe centrale et orientale, et une seconde, plus petite, dans la péninsule Ibérique.
Blarinoides, Soricidae, Lipotyphla, Pliocene, SpainBlarinoides, Soricidae, Lipotyphla, Pliocène, EspagnepresentedPresented by Yves CoppensVersion française abrégéeIntroduction
Le genre Blarinoides était jusqu’à aujourd’hui monospécifique, et connu uniquement par son espèce-type, Blarinoides mariae. Sa distribution géographique était pratiquement limitée à l’Europe centrale et orientale : Hongrie, Pologne, Bulgarie, Roumanie, Slovaquie et Allemagne [18]. Nous y ajoutons une deuxième espèce, Blarinoides aliciae, provenant du Villafranchien ancien de Tollo de Chiclana-3 (TCH-3), gisement situé dans la dépression de Guadix (Espagne sud-orientale).
Paléontologie systématique
Ordre : Lipotyphla Haeckel, 1866
Famille : Soricidae Fischer von Waldheim, 1817
Sous-famille : Soricinae Fischer von Waldheim, 1817
Tribu : Blarinini Kretzoi, 1965
Genre : Blarinoides Sulimski, 1959
Blarinoides aliciae sp. nov. (Fig. 1)
Derivatio nominis : cette espèce est dédiée à Alicia Serna Barquero, amie et collègue du Departamento de Estratigrafía y Paleontología (université de Grenade), en témoignage de reconnaissance pour son aide.
Holotype : M1 droite, TCH-3 278, conservée au Departamento de Estratigrafía y Paleontología de l’université de Grenade.
Hypodigme : 6 I inf., 8 M1, 9 M2, 5 M3, 4 I sup., 8 P4, 10 M1, 7 M2.
Localité-type : Tollo de Chiclana-3 (TCH-3), Villafranchian ancien, dépression de Guadix, Espagne. Coordonnées UTM 30SVG957549 [11].
Diagnose :Blarinoides, avec des dents morphologiquement similaires à celles de B. mariae, mais notablement plus petites. I inf. tricuspidée. M1 et M2 avec l’entoconide bien développé, et sans entocristide. M3 à hypoconide et entoconide distincts. I sup. non bifide. P4 à protocône et hypocône distincts. M1 et M2 avec une émargination postérieure faible, à métalophe long, reliant le protocône au métacône.
Diagnose différentielle :B. aliciae diffère de B. mariae principalement par sa taille inférieure. En plus, la P4 de B. aliciae présente une émargination postérieure plus forte que B. mariae. B. aliciae se distingue des espèces du genre Mafia Reumer, 1984 par sa plus grande taille, et par la présence d’un hypocône et d’un métalophe distincts sur les M1 et M2. La nouvelle espèce diffère de Sulimskia Reumer, 1984 par sa taille beaucoup plus conséquente, et par le cingulum labial de la I inf., moins développé.
La I sup. non bifide et la I inf. tricuspidée permettent de distinguer B. aliciae de toutes les espèces de Anourosoricini, Beremendiini et Neomyini. La nouvelle espèce diffère des tribus Blarinellini et Soricini par l’absence d’une entocristide en M1 et M2.
Mesures : voir Table 1 and Table 2.
Discussion et conclusions
B. mariae présente une distribution temporelle longue, du Pliocène ancien jusqu’au Pléistocène ancien, plus ancien et plus récent que TCH-3, sans présenter de variations morphologiques ou biométriques [15] and [17]. B. aliciae se distingue de B. mariae par plusieurs caractères morphologiques de la dentition, et surtout par une taille notablement plus petite (Fig. 2 and Fig. 3).
Jammot [9] a décrit une espèce, B. chalinei, qui doit être considérée comme un nomen nudum, puisque ce travail n’est pas publié.
Dans la péninsule Ibérique, Blarinoides a été cité du gisement d’Escorihuela (Villafranchien ancien) et Escorihuela B (Ruscinien tardif) [10] où le matériel est trop pauvre pour permettre une détermination. Le genre est aussi documenté par du matériel non publié de Concud Village 3 et Concud Estación 2. Au vu de la petite taille de ces matériaux, ils peuvent être attribués à B. aliciae.
Introduction
The genus Blarinoides was until now a monospecific genus, represented solely by the type species, Blarinoides mariae Sulimski, 1959. Blarinoides is one of the most common elements in the European Pliocene soricid faunas. It is quite abundant in localities from Hungary, Poland, Bulgaria, Romania, Slovakia, and Germany, ranging in age from Early Ruscinian to Early Pleistocene [18]. On the contrary, remains ascribed to Blarinoides are extremely scarce in the Iberian Peninsula, maybe due, in part, to the lack of detailed studies.
In this paper, the population of Blarinoides from the locality of Tollo de Chiclana-3 is described; this is the most numerous sample found in the Iberian Peninsula until now. The material described here shows distinctive features that allow the definition of a new species, Blarinoides aliciae sp. nov.
The section of Tollo de Chiclana is situated in the central part of the Guadix Basin (southeastern Spain), an intramontane basin established in the Late Miocene. The oldest sediments are marine Tortonian. Continental sedimentation begins in the Latest Tortonian and lasts until Late Pleistocene [5] and [21]. In the section of Tollo de Chiclana, six fossiliferous localities were described in previous publications [12] and [13]. These fossiliferous levels, ranging in age between Late Ruscinian and Middle Villafranchian, are found in swamp deposits. The faunal list of the locality of Tollo de Chiclana-3 (TCH-3) includes the following species: Stephanomys thaleri, S. minor, Castillomys crusafonti, Apodemus atavus, Mimomys stehlini, Kislangia ischus, Eliomys intermedius, Asoriculus gibberodon, Paenelimnoecus pannonicus, B. aliciae nov. sp., ‘Myosorex meini’, Archaeodesmana sp., and Talpinae indet. [11]. This association allows us to assign the locality of TCH-3 to the Early Villafranchian.
The nomenclature used in the descriptions of the teeth is that of Reumer [15]. Lengths and widths have been measured as defined by Reumer [15]. Specimens are kept in the ‘Departamento de Estratigrafía y Paleontología’ of the University of Granada, Spain.
Systematic Palaeontology
Order Lipotyphla Haeckel, 1866
Family Soricidae Fischer von Waldheim, 1817
Subfamily Soricinae Fischer von Waldheim, 1817
Tribe Blarinini Kretzoi, 1965
Genus Blarinoides Sulimski, 1959
Blarinoides aliciae sp. nov. (Fig. 1)
Derivatio nominis: this species is devoted to Alicia Serna Barquero, friend and colleague of the ‘Departamento de Estratigrafía y Paleontología’ (University of Granada), in appreciation of her help in this work.
Holotype: right M1, TCH-3 278, kept in the ‘Departamento de Estratigrafía y Paleontología’ (University of Granada).
Referred material: 6 I inf., 8 M1, 9 M2, 5 M3, 4 I sup., 8 P4, 10 M1, 7 M2.
Type locality: Tollo de Chiclana-3 (TCH-3), Early Villafranchian, Guadix Basin, Spain. Coordinates UTM 30SVG957549 [11].
Diagnosis:Blarinoides with dental elements morphologically similar to those of B. mariae, but notably smaller. I inf. tricuspulate. M1 and M2 with well-developed entoconid, and without entoconid crest. M3 with distinct hypoconid and entoconid. I sup. not fissident. P4 with distinct protocone and hypocone. M1 and M2 with a weak posterior emargination, and with a long metaloph connecting the protocone to the metacone.
Differential diagnosis:B. aliciae differs from B. mariae mainly by its smaller size. Besides, the P4 of B. aliciae has a stronger posterior emargination than B. mariae. B. aliciae can be distinguished from the species of the genus Mafia Reumer, 1984 by its larger size, and by the presence of a distinct hypocone and metaloph in M1 and M2. The new species differs from Sulimskia Reumer, 1984 by its much larger size, and by the less developed labial cingulum in the I inf.
The non-fissident I sup. and the tricuspulate I inf. allow us to distinguish B. aliciae from all the species of Anourosoricini, Beremendiini, and Neomyini. The new species differs from the tribes Blarinellini and Soricini in the lack of an entoconid crest in M1 and M2.
Measurements: see Table 1 and Table 2.
Description
I inf.: tricuspulate incisors. In unworn specimens, one may observe that the posterior cuspule is weak. The apex is notably upturned. The specimens preserving the posterior part show a weakly developed cingulum along the dorsal part of the posterolabial edge.
M1and M2: large molars, with sharp cuspids and high crests. In the M1, the trigonid is somewhat longer and notably narrower than the talonid; in the M2, trigonid and talonid have similar dimensions. The entoconid is high, and well separated from the metaconid. There is no entoconid crest. The entoconid is close to the hypolophid, separated from it by a narrow, deep valley. The labial cingulum is not very prominent, but well defined. It is low between the paraconid and the protoconid, and between the hypoconid and the entostylid; this cingulum shows an elevation at the level of the re-entrant valley. On the contrary, the lingual cingulum has a constant height. The re-entrant valley opens high on the labial cingulum. There are two roots, strongly inclined backwards.
The mental foramen is placed below the talonid of the M1.
M3: the lingual border is straight. The paraconid, protoconid and metaconid, and the crests connecting these cuspids are high. In the least-worn specimens, the entoconid and the hypoconid can be observed. The oblique crest and the hypolophid are curved, so the talonid basin is rounded. The labial cingulum is very well developed; it is more prominent in its anterior part, and it rises at the level of the re-entrant valley. The labial cingulum disappears below the hypoconid. The lingual cingulum is less prominent than the labial one, but it is similar in height and well marked. The re-entrant valley opens near the labial cingulum. There are two long and inclined roots.
I sup.: large, non-fissident incisors. The labial cingulum is slightly marked; it extends along two thirds of the posterolabial margin. The talon is weakly developed.
P4: long teeth, with trapezoidal outline. The paracone is high and sharp. The parastyle protrudes notably over the anterior border of the tooth; it is higher than the protocone and hypocone. The parastyle is connected to the paracone by a parastylar crest, and to the protocone by another crest. The protocone is large and distinct. It is connected to the hypocone by a posterolingual crest. The hypocone is well individualized, somewhat lower than the protocone. In the least-worn specimens, an extra cingular cusp can be observed behind the hypoconid. Behind this cusp, there is a short and thick posteroloph. The hypoconal flange does not protrude far backwards, and the posterior emargination is weak. Along the posterior side of the tooth, there is a narrow, but well-marked, cingulum. There are two roots with round section in anterolabial and posterolabial position, and a third one with an elongated section on the lingual side of the tooth.
M1and M2: the outline is practically quadratic in M1. The M2 is trapezoidal in occlusal view: the posterior border is shorter than the anterior one, and the labial and lingual borders converge backwards. The metacone is slightly higher than the paracone. The posterior arm of the metacone is somewhat longer than the anterior one, and the metastyle protrudes over the labial border, especially in the M1. The mesostyle is split. The protocone is lower than the paracone. There is a minuscule protoconule near the base of the paracone, separated from it by a narrow valley, and connected to the protocone by a crest. Behind the protocone, there is a long metaloph that reaches the base of the metacone, closing the trigon basin. Eight out of ten M1 have a narrow cingulum at the base of the protocone. In four specimens, the cingulum is well defined, and it extends along the anterolingual border of the molar. In the other four specimens, the cingulum is weak and short, limited to the base of the anterior side of the protocone. This cingulum only occurs in one out of seven M2. The hypocone is low and rounded, connected to the metaloph by a low anterior crest. The posteroloph is thick. There is a well-developed cingulum along the posterior border. The hypoconal flange is small; the posterior emargination is not much pronounced in the M2 and even weaker in the M1. There are four small roots with round section in anterolingual, anterolabial, posterolingual, and posterolabial positions.
No M3 is preserved in the material from TCH-3.
Discussion
Blarinoides was a monospecific genus until now. The diagnosis of the genus and species is based on dental and cranial features. Unfortunately, in TCH-3, no complete skull or mandible has been preserved. Dental elements are isolated (exceptionally two teeth are preserved in a mandible fragment), so only dental features can be observed in this population.
The dental characters of the genus Blarinoides described by Sulimski [20], which can be considered diagnostic, are: lower incisor tricuspulate, M3 with distinct hypoconid and entoconid, upper molars with a weak posterior emargination. Rzebik-Kowalska [17] gave more details on the dental morphology of B. mariae, based on several Polish localities (Wéze I, Zamskowa Dolna, and Rebielice Królewskie I and II): the upper incisor is ‘not bifid’ (not fissident, after Reumer [15]); the P4 has a large parastyle, a well-developed protocone and hypocone, and an accessory cusp behind the hypocone; the upper molars have a metaloph; the occlusal outline of the M2 is trapezoidal; the lower molars have a large entoconid, but lack an entoconid crest. All these features are found in other populations of B. mariae, described in the literature (like Csarnóta 2, Osztramos 7 and 9 [15]), and can also be observed in the population from TCH-3.
We have compared the material from TCH-3 with that from the type locality of B. mariae, Wéze, stored at the University of Lyon-1 (France). The most important morphological differences are: in the M1 of B. mariae from Wéze, the difference in width between the trigonid and the talonid is subtle, and the labial border is almost straight and parallel to the lingual one. On the contrary, in the specimens of B. aliciae from TCH-3, the trigonid is markedly narrower than the talonid; besides, on the occlusal outline, there is a concavity at the level of the re-entrant valley, which does not exist in the specimens from Wéze. The labial cingulum is somewhat higher in the M1–3 of B. mariae than in those of B. aliciae. The P4 from Wéze have a notably weaker posterior emargination than those from TCH-3; this character can be also observed in the specimens from other populations of B. mariae (Osztramos 7, Csarnóta 2), figured by Reumer [15].
Biometrical differences between all the material assigned to B. mariae and the population from TCH-3 are much more evident. Sulimski [20] gives no measurements of isolated teeth. Rzebik-Kowalska [17] published measurements of the upper dentition only, which are based on a measurement method [16] different from ours. Nevertheless, it is clear that the size of B. mariae from the localities of Wéze I, Zamskowa Dolna, and Rebielice Królewskie I and II is notably larger than that of B. aliciae from TCH-3. Direct comparison of the material from TCH-3 with that from Wéze has allowed us to observe a notable difference in size for all dental elements, especially for the incisors, which are much larger in B. mariae from Wéze.
The size of the specimens from Csarnóta 2, Osztramos 7 and 9 [15] is also clearly larger than that of B. aliciae from TCH-3, for all the measured parameters. The maximum values in TCH-3 are usually smaller than the minimum values in these Hungarian populations. For some dental elements, especially the M3 and the M2, certain overlap of the size ranges exists between the population of TCH-3 and those of Csarnóta and Osztramos. Anyway, the mean values are clearly smaller in the population from TCH-3 for all the considered parameters. The size of B. aliciae from TCH-3 is also smaller than that of the few specimens of B. mariae found in Betfia-XIII (Romania, [19]), Muselievo (Bulgaria, [14]), and Vue-des-Alpes (Switzerland, [2]). Fig. 2 and Fig. 3 represent the ranges of variation of several parameters of the dentition of B. aliciae from TCH-3 and B. mariae from various European localities.
It is important to emphasize that we have compared TCH-3 with populations of B. mariae both older (Osztramos 9, Csarnóta 2, Muselievo, Vue-des-Alpes), and younger (Osztramos 7, Betfia-XIII) than it; the sizes of all of them are notably larger than that of B. aliciae. B. mariae has a long temporal distribution (from Early Pliocene to Early Pleistocene); during this time interval, no morphological or biometrical changes can be observed, as Rzebik-Kowalska [17] and Reumer [15] pointed out.
In the Iberian Peninsula, B. mariae has been cited from the locality of Escorihuela (Lower Villafranchian) and Blarinoides sp. from Escorihuela B (Upper Ruscinian) [10]. The material from TCH-3 has been compared with that from these localities, and also with unpublished material from Concud Village 3 and Concud Estación 2, stored at the University of Lyon-1. The material from Escorihuela and Escorihuela B is very scarce (a single M1 in the former locality, one M1 and one M2 in the latter). These specimens are slightly larger than those of B. aliciae from TCH-3. Nevertheless, the scarcity of the material does not allow us to do a specific determination, so we refer to it as Blarinoides sp. The material from Concud Village 3 and Concud Estación 2 is more abundant. The specimens from these localities are similar in size to those from TCH-3, and notably smaller than those of B. mariae from Wéze, and can be assigned to the new species B. aliciae. Therefore, the geographical distribution of this species included the greater part of the Iberian Peninsula.
Crochet [4] cited for the first time the possible existence of two different lineages of the genus Blarinoides in Europe. He assigned a single M2 from Terrats (Roussillon), morphologically similar to B. mariae, but notably smaller, to Blarinoides sp. This specimen is clearly smaller than those from TCH-3, and it is morphologically similar to the specimens of B. aliciae, but its hypocone is less developed. The locality of Terrats (Early Pliocene, MN14) is much older than TCH-3, Concud Village 3, and Concud Estación 2. If the generic determination of the single molar from Terrats were correct, this would indicate the presence of a small-sized lineage of Blarinoides in western Europe since the beginning of the Pliocene. However, its adscription to this genus is uncertain, because of the scarcity of material and of the pronounced difference in size with all known populations of Blarinoides. More numerous samples from this locality would be necessary to determine this specimen beyond doubt and clarify its possible relation with B. aliciae. For the moment, the single tooth from Terrats cannot be determined at a specific level; therefore, we keep the denomination of Blarinoides sp.
Jammot [9] described a new species of this genus, Blarinoides chalinei, from the locality of Seynes. This paper is unpublished, and hence this species is considered a nomen nudum. However, we have revised the manuscript of Jammot, in order to check the possible resemblance between the populations from TCH-3 and Seynes. The diagnosis of Jammot is based mainly on cranial features. The description of the teeth is brief and not detailed: the diagnosis only specifies that the lower molars have a well-developed labial cingulum, there is no reference to the morphology of the upper molars, and there are no tooth measurements. Jammot does not mention the difference in size between ‘B. chalinei’ and B. mariae, and indicates that the difference between these two species is based on the shape of the mandible. Therefore, it seems very unlikely that the population from Seynes would correspond to B. aliciae, which differs notably in size from B. mariae.
The presence of Blarinoides has been mentioned in other few western European localities: B. mariae from Hautimagne (France) [6] and from Arondelli (Italy) [1], and Blarinoides sp. from Arcille (Italy) [8]. It would be very interesting to analyze the size and the morphology of these populations to ensure their specific ascription and to clarify their relationships with the species B. mariae and B. aliciae. Nevertheless, the material from these localities is very scarce and there are no published descriptions, measurements or figures of the specimens; therefore, it is impossible to interpret their relationships with other European populations. Table 3 compiles the current available data of the presence of Blarinoides in Europe.
B. aliciae from TCH-3 differs clearly from the rest of the Soricidae found in the Pliocene of Europe. It is larger than Crocidura, and very different morphologically: the upper molars of B. aliciae are longer, with a weaker posterior emargination, and the lower molars have a less compressed talonid and a deeper re-entrant valley. The new species differs from Paenelimnoecus in its much larger size, and in the presence of an entoconid in the lower molars. With regard to the Soricinae, B. aliciae can be distinguished from all the Soricini, Neomyini, Blarinellini, and Beremendiini in two characters: non-fissident upper incisor and lack of an entoconid crest in the lower molars. B. aliciae differs from the Anourosoricini in the tricuspulate lower incisor and in the well-developed talonid basin in the M3. Furthermore, B. aliciae is much larger than most of the fossil Soricidae. Its size is only comparable to that of Beremendia, but this genus differs obviously from Blarinoides in the fissident upper incisor, the acuspulate lower incisor, the presence of entoconid crests in the lower molars, and the stronger posterior emargination in the upper molars.
B. aliciae can be distinguished from the other genera of the tribe Blarinini found in the Pliocene of Europe, Sulimskia and Mafia, mainly by its larger size. Besides, the new species has some distinctive morphological features, like the labial cingulum of the lower incisor, which is weaker than in Sulimskia, and the hypocone in the upper molars, more developed than in Mafia.
Conclusion
The population of B. aliciae sp. nov. from the locality of TCH-3 is the most numerous population of Blarinoides found in the Iberian Peninsula until now, and represents the first citation of the genus in southern Spain. This new species is morphologically very similar to B. mariae, but markedly smaller. Apart from the type population, the material from Concud Village 3 and Concud Estación 2 is assigned to this species.
Blarinoides now comprises two contemporaneous species: B. mariae, from central and eastern Europe, and B. aliciae, from the Iberian Peninsula.
Acknowledgements
This study was supported by the projects CGL2006–4548/BTE (Spanish Ministry of Education and Science) and Programa Consolider-Ingenio 2010 (CSD2006–00041), and the group ‘Paleogeografía de Cuencas Sedimentarias’ (RNM190) of the ‘Junta de Andalucía’. The photographs were made using the Zeiss 950 Scanning Electron Microscope of the ‘Centro de Instrumentación Científica’ of the University of Granada. We thank Pierre Mein, who put the material stored at the University of Lyon-1 at our disposal. Comments and suggestions by anonymous reviewers are kindly acknowledged.
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Blarinoides aliciae sp. nov. from TCH-3, Guadix Basin. A, Right I inf., TCH-3 289; B, right I sup., TCH-3 262; C, right M3, TCH-3 259; D, right M2, TCH-3 253; E, right M1, TCH-3 243; F, left M1, TCH-3 241, labial view; G, left M2, TCH-3 247, labial view; H, left M3, TCH-3 256, labial view; I, right M2, TCH-3 285; J, right M1, TCH-3 278 (holotype); K, right P4, TCH-3 269. Scale = 1 mm.
Ranges of variation of several parameters of the dentition of Blarinoides mariae from Osztramos 7 and 9, Csarnóta 2 [15], and Betfia XIII [19], and of Blarinoides aliciae from TCH-3. A, length of the I inf.; B, length of the I sup.; C, length of the M1; D, talonid width of the M1; E, length of the M2; F, talonid width of the M2.
Fig. 2. Variation de quelques paramètres de la dentition de Blarinoides mariae de Osztramos 7 et 9, Csarnóta 2 [15] et Betfia XIII [19], et de Blarinoides aliciae de TCH-3. A, longueur de I inf. ; B, longueur de I sup. ; C, longueur de M1 ; D, largeur du talonide de M1 ; E, longueur de M2 ; F, largeur du talonide de M2.
Ranges of variation of several parameters of the dentition of Blarinoides mariae from Osztramos 7 and 9, Csarnóta 2 [15], and Betfia XIII [19], and of Blarinoides aliciae from TCH-3. A, buccal length of the P4; B, width of the P4; C, buccal length of the M1; D, anterior width of the M1; E, buccal length of the M2; F, anterior width of the M2.
Fig. 3. Variation de quelques paramètres de la dentition de Blarinoides mariae de Osztramos 7 et 9, Csarnóta 2 [15], et Betfia XIII [19], et de Blarinoides aliciae de TCH-3. A, longueur buccale de P4 ; B, largeur de P4 ; C, longueur buccale de M1 ; D, largeur antérieure de M1 ; E, longueur buccale de M2 ; F, largeur antérieure de M2.
Measurements (mm) of the lower teeth of Blarinoides aliciae sp. nov. from Tollo de Chiclana-3
Tableau 1 Mesures (en mm) des dents inférieures de Blarinoides aliciae sp. nov. de Tollo de Chiclana-3