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Historical BiologyAn International Journal of Paleobiology

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A new species of Plesiodimylus (Dimylidae,Eulipotyphla, Mammalia) from the Early Mioceneof Spain

Vicente D. Crespo, Marc Furió, Francisco Javier Ruiz-Sánchez & Plini Montoya

To cite this article: Vicente D. Crespo, Marc Furió, Francisco Javier Ruiz-Sánchez & Plini Montoya(2017): A new species of Plesiodimylus (Dimylidae, Eulipotyphla, Mammalia) from the EarlyMiocene of Spain, Historical Biology

To link to this article: http://dx.doi.org/10.1080/08912963.2017.1289519

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Historical Biology, 2017http://dx.doi.org/10.1080/08912963.2017.1289519

A new species of Plesiodimylus (Dimylidae, Eulipotyphla, Mammalia) from the Early Miocene of Spain

Vicente D. Crespoa,b, Marc Furióc, Francisco Javier Ruiz-Sáncheza,b,d and Plini Montoyaa

aDepartament de Botànica i geologia, Universitat de València, Valencia, spain; bMuseu Valencià d’Història Natural, l’Hort de Feliu, Valencia, spain; cinstitut català de Paleontologia M. crusafont, Edifici Z (icta-icP) c/ de les columnes s/n, Universitat autònoma de Barcelona, Barcelona, spain; diNcyt-UPsE, Universidad Peninsula de santa Elena, santa Elena, Ecuador

ABSTRACTWe report a new dimylid species, Plesiodimylus ilercavonicus sp. nov., from the Early Miocene locality of Mas d’Antolino B-5 (Ribesalbes-Alcora, Castelló, Spain). This new species of Plesiodimylus is an amblyodont form of the genus and exhibits some primitive characters. The phylogenetic and palaeoenvironmental implications of this southern occurrence of Plesiodimylus in Lower Miocene sediments are discussed.

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KEYWORDSaragonian; insectivores; soricomorpha; iberian Peninsula; ribesalbes-alcora Basin

ARTICLE HISTORYreceived 20 october 2016 accepted 28 January 2017

CONTACT Marc Furió marc.furio@icp.cat

Introduction

The dimylids were a rather common group of eulipotyphlans in Europe from the Late Oligocene till the Late Miocene. They dis-played peculiar dentitions (sometimes described as ‘aberrant’), which have been interpreted as adaptations to a much specialized durophagous diet (Ziegler 1999; Fejfar & Sabol 2009; Furió et al. 2011a). The unusual dental traits of this family included amblyo-donty (inflated teeth), exodaenodonty (teeth overlapping dentary in occlusal view) and dimyly (evolutionary loss of third upper and lower molars). Hitherto, there is no thorough study on the functional meaning for such an extreme modified dentition, but it is commonly accepted that these bulbous teeth were likely an adaptation to a malacophagous diet (Müller 1967).

The heydays of the family correspond to the Early Miocene, reaching a maximum in diversity in Central Europe during this time (Ziegler 1999). However, in the southern zone of Europe they were less diverse (Doukas 2003; Furió et al. 2011a). The Iberian Peninsula is a good example of low diversity, as only three gen-era of dimylids (Chainodus, Metacordylodon and Plesiodimylus) have been reported, usually restricted to few finds concentrated in some specific basins. For instance, Chainodus cf. sulcatus has only been reported in the lower Miocene from Rubielos de Mora (Teruel, MN 3; Van den Hoek Ostende & Fejfar 2015) [originally Chainodus intercedens Müller 1967 in Gibert (1975) and Montoya et al. (1996)] and the Vallès-Penedès Basin (unpublished material from Turó de les Forques, an Early Miocene site in Furió et al. 2011a). Similarly, Metacordylodon schlosseri (Andreae, 1904) has only been documented in Spain at Castell de Barberà and Sant Quirze A (Crusafont-Pairó & Golpe 1972; Van den Hoek Ostende & Furió 2005; Prieto et al. research in progress).

The genus Plesiodimylus was the most frequent representative of the family in the Miocene, ranging from the late Early Miocene

to the early Late Miocene (Ziegler 1999). Its occurrences in Spain had been hitherto limited to only one species, Plesiodimylus chant-rei Gaillard, 1899 (Furió et al. 2011b), being rather common in the vertebrate assemblages of the Vallès-Penedès Basin during the late Aragonian and the Vallesian (Crusafont & Truyols 1951; Van den Hoek Ostende & Furió 2005; 2011b; Furió et al. 2011a; Casanovas-Vilar et al. 2012). The genus had never been reported further to the south in Spain. Its limited temporal and geographical range in the Iberian Peninsula led to the hypothesis that Plesiodimylus, though displaying the less specialised dental features within the Dimylidae, had a diet too much specialized to reach a wider dis-tribution in both time and space (Furió et al. 2011a, b).

Field works carried out in 2008 and 2010 in the locality of Mas d’Antolino B-5 (Ribesalbes-Alcora Basin, Castelló province, Figure 1) have provided new fossils of Plesiodimylus. They rep-resent the first record of the genus in Lower Miocene sediments from Spain. Moreover, this site is the southernmost occurrence ever reported for Plesiodimylus. Paradoxically, the dental features of this species were more amblyodont than in its Middle-Late Miocene counterparts from the Vallès-Penedès, thus opening the question to what extent the distribution of Plesiodimylus was constrained during the Miocene by ecological reasons, and if diet was indeed the limiting factor. In order to elucidate the palae-obiological profile of this genus and its ecological preferences, we describe and illustrate the dental features of this new species and its wear pattern.

Geographic and Geologic Setting

The fossil material described in the present work comes from the western area of the Ribesalbes-Alcora Basin (Figure 1), an intramontane basin in eastern Spain (Agustí et al. 1988). These

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2 V. D. CRESPO ET AL.

Methods. This paper follows the nomenclature and measure-ments used by Müller (1967) updated in Klietmann et al. (2014). Measurements (L x W) are given in millimetres and were taken using a Leica MZ75 binocular microscope, by means of displace-ment of a mechanical stage, connected to a Sony Magnescale measuring equipment. The lower teeth are written in lower-case letters (lower molars: m1 and m2; lower premolars: p1, p2, p3 and p4; lower canine: c) and the upper teeth are written in upper-case letters (upper molars: M1 and M2; upper premolars: P1, P2, P3, and P4; upper canine: C; upper incisors: I1, I2 and I3).

Abbreviations. Institutional. MGUV: Museu de Geologia de la Universitat de València, Burjassot (Spain); SCSIE: Servei Central de Suport a la Investigació Experimental, Valencia (Spain); UV: Universitat de València, Valencia (Spain). Localities. MAB: Mas d’Antolino B.

Systematic paleontology

Order EULIPOTYPHLA Wadell, Okada and Hasegawa, 1999Family DIMYLIDAE Schlosser, 1887Genus PLESIODIMYLUS Gaillard, 1897PLESIODIMYLUS ILERCAVONICUS SP. NOV.(Figures 2, 3, 4, 5, 6; Tables 1, 2)

Holotype: MAB5–50, right M1 (MGUV–23387, Figure 2.5).

Paratypes: right C (MAB5–416, MGUV–25206; Figure 2.1); left P1 (MAB5–477, MGUV–26752); right P2 (MAB5–474, MGUV–26749; Figure 2.2); left P3 (MAB5–422, MGUV–25212; Figure 2.3); right P4 (MAB5–454, MGUV–25244; Figure 2.4); right M1

sections are made up of about 60 meters of grey and yellow mud-stones, sandstones, and limestones, and they all belong to ‘Unit Three’ sensu Anadón (1983).

The first fossils of small mammals found in the Ribesalbes-Alcora Basin were described by Agustí et al. (1988). In the last years, several new mammal sites have been found in this basin, chiefly from the sections of Mas d’Antolino B (MAB), Barranc de Campisano (BC) and Mas de Torner (MTR) (Ruiz-Sánchez et al. 2010; Furió et al. 2010a; 2010b, 2012; Crespo et al. 2012a; 2012b). It is worth noting that this basin has delivered the south-ernmost occurrence of the herpethoteriid Amphiperatherium fre-quens (von Meyer, 1846) and the youngest record of the glirid Altomiramys (Crespo et al. 2012b; Furió et al. 2012). Within the twenty fossiliferous levels in these sections providing small mammal remains, only one site has delivered elements of Plesiodimylus: Mas d’Antolino B-5 (MAB5, a bed of dark mud-stones with plant remains and gypsum).

According to Ruiz-Sánchez et al. (2010), the preliminary study of the small mammal assemblage from MAB-5 resulted in the identification of Megacricetodon cf. primitivus Mein and Freudenthal, 1971, and Ligerimys ellipticus Daams, 1976, which are indicative of an Early Miocene age (Lower Aragonian, MN4, biozone C, ca. 16.6–16.0 Ma, according to Van der Meulen et al. 2011; 2012).

Material, methods and abbreviations

Material. Photographs were made with a SEM HITACHI 4800 at the SCSIE of the UV. The fossil material is stored at the MGUV.

Figure 1. cenozoic basins from spain, with location of the ribesalbes-alcora Basin and the schematic distribution of sediments. right margin: general schematic section of the Mas d’antolino B section with all the levels providing vertebrate fossils. the placement of the locality of Mas d’antolino B 5 in the basin and in the stratigraphic section is signaled with an asterisk (*).

HISTORICAL BIOLOGY 3

(MAB5–384, MGUV–25174; Figure 2.6), right M2 (MAB5–49, MGUV–23386; Figure 2.7); left c (MAB5–472, MGUV–26747; Figures 3.1, 5.1); right p1 (MAB5–467, MGUV–26742; Figure 3.2); right p2 (MAB5–160, MGUV–24950; Figure 3.3); left p3 (MAB5–476, MGUV–26751; Figure 3.4); left p4 (MAB5–456, MGUV–25246; Figure 3.5); left m1 (MAB5–462, MGUV–26737; Figure 3.6); left m2 (MAB5–450, MGUV–25240; Figure 3.7).

Referred material: C: MAB5–167 (MGUV–24957), MAB5–414 (MGUV–25204), MAB5–451 (MGUV–25241), MAB5–457 (MGUV–25247), MAB5–752 (MGUV–27027); P1: MAB5–166 (MGUV–24956); P2: MAB5–164 (MGUV–24954), MAB5–482 (MGUV–26757), MAB5–899 (MGUV–27174); P3: MAB5–40 (MGUV–23377), MAB5–151 (MGUV–24941), MAB5–155 (MGUV–24945), MAB5–157 (MGUV–24947), MAB5–452 (MGUV–25242), MAB5–478 (MGUV–26753), MAB5–895 (MGUV–27170), MAB5–897 (MGUV–27172), MAB5–898 (MGUV–27173); P4: MAB5–406 (MGUV–25196), MAB5–413 (MGUV–25203), MAB5–418 (MGUV–25208), MAB5–449 (MGUV–25239), MAB5–461 (MGUV–26736); M1: MAB5–48 (MGUV–23385), MAB5–50 (MGUV–23387), MAB5–150 (MGUV–24940), MAB5–385 (MGUV–25175), MAB5–392 (MGUV–25182), MAB5–418 (MGUV–25208), MAB5–448 (MGUV–25238), MAB5–453 (MGUV–25243), MAB5–459 (MGUV–26734), MAB5–470 (MGUV–26745); M2: MAB5–460 (MGUV–26735), MAB5–486 (MGUV–26761); Maxillar: MAB5–383 (with P4 alveol and M1 tooth, MGUV–25173); c: MAB5–473 (MGUV–26748), MAB5–480 (MGUV–26755); p1: MAB5–468 (MGUV–26743), MAB5–471 (MGUV–26746); p2: MAB5–159 (MGUV–24949), MAB5–421 (MGUV–25211), MAB5–896 (MGUV–27171); p3: MAB5–901 (MGUV–27176); p4: MAB5–466 (MGUV–26741), MAB5–469 (MGUV–26744); m1: MAB5–41 (MGUV–23378), MAB5–147 (MGUV–24937), MAB5–369 (MGUV–25159), MAB5–373 (MGUV–25163), MAB5–374 (MGUV–25164), MAB5–410 (MGUV–25200), MAB5–444 (MGUV–25234), MAB5–464 (MGUV–26739), MAB5–484 (MGUV–26759), MAB5–485 (MGUV–26760); m2: MAB5–143 (MGUV–24933), MAB5–375 (MGUV–25165), MAB5–446 (MGUV–25236), MAB5–463 (MGUV–26738), MAB5–465 (MGUV–26740); Mandibles: MAB5–455 (with alve-oli of i3–p4; MGUV–25245), MAB5–475 (with m2 and alveoli of m1; MGUV–26750), MAB5–722 (with alveoli of p1–p4; MGUV–26997); MAB5–723 (with p4 and alveoli of p1–p3; MGUV–26998).

Other material of difficult identification: I1, 2?: MAB5–38 (MGUV–23375), MAB5–152 (MGUV–24942), MAB5–153 (MGUV–24943), MAB5–424 (MGUV–25214), MAB5–429 (MGUV–25219), MAB5–433 (MGUV–25223), MAB5–458 (MGUV–25248); I3?: MAB5–154 (MGUV–24944), MAB5–162 (MGUV–24952), MAB5–766 (MGUV-27041); MAB5–900 (MGUV–27175).

Etymology: From ‘Ilercavons’, a pre-Roman tribe of Iberian peo-ple who lived in an area currently included in the provinces of Castelló and Tarragona (East of Spain), where the fossils herein described come from.

Diagnosis: Intermediate to large-sized species of Plesiodimylus with rather stout and amblyodont teeth within the genus; P4 almost as wide and long; M2 less reduced respect to the M1

than usual in the genus and with a complex metacone preserving several cuspules; p4 with wrinkled enamel, quadratic in occlu-sal view and well-developed cingulid; posterior cingulid in m1 obliquely ascending, reaching the postcristid.

Differential Diagnosis

Plesiodimylus ilercavonicus sp. nov. differs from P. chantrei in having a larger size, an ascending oblique cingulid at the poste-rior face of m1, a variable presence of mesostyle in M1, a divided mesostyle in M2 and a cuspulated metacone in M2.

Plesiodimylus ilercavonicus sp. nov. differs from P. gaillardi Mein & Gingsburg 2002 in its larger size, an ascending oblique cingulid at the posterior face of m1, and preserving four lower premolars.

Plesiodimylus ilercavonicus sp. nov. differs from P. bavaricus Schötz 1985 in an overall larger size, having a wider paracone of the P4, not having invariably present a mesostyle in the M1, a less developed posterior cingulum in M1, a rounded proto-cone and the anterior arm not reaching the paracone in M1, a sharp metastyle and in having a more complex structure of the metacone in M2.

Plesiodimylus ilercavonicus sp. nov. differs from P. similis Fejfar & Sabol 2009 in its somewhat smaller teeth (except upper and lower second molars, which are comparable in size), the longer trigonids and narrower talonids of its lower molars, having a little cusp close to the paraconid of the m2, having a P4 with a less developed cingulum, a more pronounced parastyle in the M1, and in having a more irregular labial margin of the M1 in occlusal view.

Plesiodimylus ilercavonicus sp. nov. differs from P. huerzeleri Müller 1967 in having a larger size, the variable presence of meso-style, constant presence of cuspular parastyle, an anterior arm of parastyle reaching the paracone, and having three basins in M1, displaying a more complex metacone in M2 with several cus-pules, and having a relatively longer M2 with respect to the M1.

Plesiodimylus ilercavonicus sp. nov. differs from P. helveticus Bolliger 1992 in its lower molars with wider talonids, having a less pronounced reentrant valley in m1, the more rounded trigonid of the m2, showing a wider anterior basal cingulid of the m2, showing a P4 with a more concave anterolingual side in occlusal view and a more advanced protocone, having a cuspular parastyle in M1 and in having a M2 with a conical protocone and a more complex metacone.

Plesiodimylus ilercavonicus sp. nov. differs from P. johanni Kälin & Engesser 2001 in having a shorter p4 and a wider P4, a M1 with distinct parastyle combined with a variable presence of the mesostyle, and in its shorter postparacrista in this tooth.

Plesiodimylus ilercavonicus sp. nov. differs from P. crassi-dens Engesser 1980 in its smaller size, having a rather squarish occlusal outline of the p4, the more irregular labial margin of the M1 in occlusal view, and in the more complex metacone in M2.

Plesiodimylus ilercavonicus sp. nov. differs from Dimylus para-doxus von Meyer, 1846 in having (on average) m2 longer than the m1, having a protoconid and metaconid standing closer to each other and not having entostylid in m1, and in a more developed protocone of the P4.

4 V. D. CRESPO ET AL.

Synonymy

2010-Dimylidae indet., in Furió et al. 2010a, p. 126.2010-Plesiodimylus sp., in Furió et al. 2010b, p. 93A.2012-Plesiodimylus sp., in Furió et al. 2012, p.374, Tab. 2.Type Locality: Mas d’Antolino B-5, Araia d’Alcora, Alcora, Castelló Province, East of Spain.

Stratigraphic correlation: Lower Miocene (lower Aragonian, MN4) from Spain. 16.6–16.0 Ma.

Measurements: see Table 1

Description of the material

Maxillary

One of the maxillary fragments found preserves a small part of the zygomatic arch and the M1. The zygomatic arch is broken. The infraorbital foramen is elliptical in frontal view and placed above the position of P4. The lachrymal foramen is rather cir-cular and it is located dorsocaudal to the infraorbital foramen, above the contact between P4 and M1.

Upper Dentition

The C (Figure 2.1) is high crowned and tear-shaped in occlu-sal view. There is a continuous basal cingulum surrounding the entire tooth. There are two crests, a high posterior one from top of the tooth to the cingulum, and a faint anterolingual one. The tooth is double-rooted with parallel roots. According to the specimens available, as the wear advances, it creates a flat surface progressively larger at the posterolingual side of the tooth.

The P1 has a laterally compressed oval occlusal outline. Its basal cingulum is poorly developed. The main cusp is high, and it occupies an anterior position. A crest crosses the tooth antero-posteriorly.

The P2 (Figure 2.2) is the smallest upper tooth, and it has a sub-circular occlusal outline. There is a well-developed cingulum covering the labial side of the tooth. There is only paracone in the middle of an unconspicuous longitudinal crest. The anterior part of the crest is shorter than the posterior one, so the paracone occupies an antero-central position.

The P3 (Figure 2.3) is somewhat larger than the P2. It has a subquadrate occlusal outline with rounded corners. The para-cone has an antero-central position. The cingulum covers com-pletely the base of the crown. In four out of seven P3, there is an expansion of this cingulum at the postero-lingual side bearing a little cuspule. The paracone occupies a middle-to-anterior posi-tion. The anterior crest is not as sharp as the posterior crest. Two of the three specimens preserving roots show two fused roots, whereas the third specimen shows an anterior root but its posterior one is broken.

The P4 (Figure 2.4) is subtriangular with rounded margins in occlusal view. The width of the P4 is almost similar to its length in most specimens. The position of the parastyle is occupied by a small elevation of the basal cingulum, but it is only present in three out of five specimens. The paracone is high and stout, with a ridge on its posterior slope connecting to the basal cingulum. The protocone is conical and completely isolated from the paracone.

Table 1.  Measurements of the teeth of Plesiodimylus ilercavonicus nov. sp. from MaB5.

c l WMgUV–24957 1.31 –MgUV–25204 1.43 0.89MgUV–25206 1.43 0.88MgUV–25241 – 0.87MgUV–25247 – 0.81MgUV-27027 1.39 0.87P2 l WMgUV–24954 – –MgUV–26749 0.83 0.66MgUV–26754 0.74 0.63MgUV–26757 0.78 0.65MgUV–27174 – –P3 l WMgUV–23377 0.96 0.77MgUV–24941 0.92 0.85*MgUV–24945 1.01 0.82MgUV–24947 0.88 0.75MgUV–25212 0.96 0.82MgUV–25242 0.88 0.76MgUV–26753 0.89 0.86MgUV–27169 – 0.84MgUV–27170 0.83 –MgUV–27172 0.91 –MgUV–27173 – –P4 l WMgUV-25195 1.96 1.83MgUV-25196 – –MgUV-26736 1.90 1.87*MgUV-25244 2.04 1.80MgUV-25239 2.06 1.76MgUV-25203 – –MgUV-23375 – –M1 l WMgUV-24940 – –MgUV-25238 – –MgUV-25243 – 2.31MgUV-26734 3.30 2.29MgUV-26745 – –MgUV-25208 – –MgUV-25173 3.28 2.47*MgUV-25174 3.27 2.27MgUV-25175 – 2.38MgUV-25182 – –MgUV-23385 – 2.14*MgUV-23387 – 2.48MgUV-25209 – –M2 l WMgUV-23386 2.14 2.77* MgUV-26735 2.20 2.89MgUV-26761 – –MgUV-26638 – –MgUV-26639 – –MgUV-26637 2.11 –MgUV-27188 – –c l WMgUV–26747 – 0.84MgUV–26748 1.24 0.91MgUV–26755 1.29 0.91p1 l W*MgUV–26742 1.74 1.11MgUV–26743 1.75 1.12MgUV–26746 1.63 1.04p2 l WMgUV–24949 0.70 0.63*MgUV–24950 0.69 0.73MgUV–27171 0.62 0.68MgUV–27181 – 0.72p3 l WMgUV–26751 0.78 0.77MgUV–27176 0.79 –p4 l W*MgUV-25246 1.52 1.02MgUV-26741 1.48 1.08MgUV-26744 1.51 1.05

(Continued)

HISTORICAL BIOLOGY 5

as a slight protruding posterolabial edge of the postmetacrista. In all the specimens the posterior margin is convex in occlusal view and it bears a marked cingulum, which closes the posterior valley. The anterior base of the protocone is connected to the parastyle by a tiny and short ridge. The postprotocrista runs from the protocone to the anterior base of the hypocone. There is a slight ‘central’ elevation labially to the most posterior extreme of the postprotocrista. This elevation is well developed in two out of eight, and double in one specimen. The hypocone is rather conical and similar in size to the protocone. The basal cingulum is only interrupted at the anterior border and below the poster-olingual margin of the hypocone.

The M2 (Figure 2.7) is subtriangular in occlusal view. Both anterior and posterior cingula connect with the parastyle. The anterior cingulum is not conected to the protocone. The pro-tocone is a low conical cusp isolated from any other element of the tooth. The paracone is only somewhat higher than the protocone. Postparacrista and premetacrista meet in a slightly divided mesostyle. The metacone is complex and divided in three little elevations. There is a small notch between the lingual and central cusp. The complexity of the metacone is overdeveloped in one specimen (MAB5–49).

Mandible

At least one mental foramen is present under the m1, placed between its two roots. The internal temporal fossa extends ante-riorly till below the most posterior part of the m2.

Lower Dentition

The c (Figure 3.1) is triangular in occlusal outline, with a pos-terior margin shorter than the two labial sides. The main cusp is placed in a much anterior position. The cingulid covers most

The protocone is placed lingually to the paracone, always exactly at the same longitudinal position. The wear drastically affects the paracone creating a flat horizontal surface. The wear creates a similar effect on the protocone, but it starts somewhat before the flat surface of the paracone reaches the height of the proto-cone. Thanks to this flat wear surface, it is possible to see that the enamel layer is thick. The less worn specimens show that the basal cingulum is only interrupted at the anterolingual side of the protocone.

The M1 (Figures 2.5, 2.6) has an ‘inverse-trapezoidal’ outline in occlusal view. It has a small parastyle and a compressed para-cone. The mesostyle is visible as a small cusp developed labially to the posterior part of the postparacrista in two out of eight specimens. The mesostyle is only slightly developed in five of them, and it is completely absent in the eighth specimen. The metacone is the highest and most prominent cusp of the M1. The metastyle is not represented as a cusp, but it is only discernible

Figure 2.  Upper teeth in occlusal view of Plesiodimylus ilercavonicus sp. nov. from MaB5, type material. 1- right c (reversed, MgUV–25206; Paratype); 2- right P2 (reversed, MgUV–26749; Paratype); 3- left P3 (MgUV–25212; Paratype); 4- right P4 (reversed, MgUV–25244; Paratype); 5- right M1 (reversed, MgUV–23387; Holotype); 6- right M1 (reversed, MgUV–25174; Paratype); 7- right M2 (reversed, MgUV–23386; Paratype); 8- left i1/2? (MgUV–25248); 9- left i1/2? (MgUV-25223); 10- left i3? (MgUV-24952).

Table 1. (Continued)

type material is indicated with *.

MgUV-26998 1.55 1.06m1 l Wtr WtaMgUV-25163 – – –MgUV-25164 – – –MgUV-25234 – – 1.41MgUV-26739 2.47 1.41 1.57MgUV-25159 2.64 1.44 1.48*MgUV-26737 2.54 1.36 1.48MgUV-23378 – – 1.42MgUV-26759 – – 1.57m2 l W*MgUV-25240 2.80 1.46MgUV-27019 – 1.52MgUV-25165 2.61 –MgUV-26740 – –MgUV-26738 2.75 1.48MgUV-26760 – –MgUV-25236 – –MgUV-26750 2.70 1.50

6 V. D. CRESPO ET AL.

the hypoconid to the entoconid, interrupting the postcristid at its lingual half.

The m2 (Figure 3.7) has a trigonid wider than the talonid. The paraconid is rather high and there is a tiny lingual cusp associated to it by a small crest. There is a posterior cingulid to this supplementary cusp that partially covers the base of the trigonid basin. The trigonid basin has a little pit near to the base of the protoconid. Paraconid and protoconid are connected by a V-shaped paracristid. The protoconid and the metaconid are not as closer as they are in m1, but both are also connected by means of a straight protolophid. In the talonid, the entoconid is higher than the hypoconid. The hypoconid is connected to the base of protoconid by means of a crest. The talonid basin is open by its anterolingual side. The labial cingulid starts in an intermediate point of the posterior side and it reaches the lingual side of the trigonid basin. The wear affects the highest parts of the trigonid and talonid, creating flat horizontal surfaces at different levels in each part.

Teeth of difficult identification

While reviewing specific literature, we discovered that the ante-rior dentition of Plesiodimylus is not much known and scarcely described. Only some schematic drawings of upper incisors were figured in Müller (1967) and Bolliger (1992). As our material includes only isolated teeth and there is no mandible or max-illary with their dental elements ‘in situ’, our identifications are tentative and the teeth could indeed have a different position.

Unknown – I1?/I2? (Figures 2.8, 2.9) The crown is oval, in occlusal view. The labial cingulum extends until the anterior border of the incisive, less in mesial side. There is only one root present. The root displays a mesial groove.

I3? (Figure 2.10) The morphology of this tooh is similar to I1, 2 but it is smaller and there is no lateral groove in its root.

Discussion

The real phylogenetic relationships between the different species of Plesiodimylus are still far from being solved. According to

of the base of the tooth, except its anterior side. There is a faint longitudinal ridge in the middle of the posterior cingulid.

The p1 (Figure 3.2) is subrectangular, with the protoconid relatively blunt and placed anteriorly. The cingulid is well devel-oped all around the base of the crown. There is a longitudinal crest crossing the tooth, from the anterior to the posterior side. There is a sulcus lingually to the posterior part of the longitudi-nal crest. In one of the three specimens available, there is a tiny elevation on the anterior side. There are two roots, being the anterior smaller than the posterior one.

The p2 (Figure 3.3) is an oval single-rooted tooth. It is the smallest lower premolar. It has a blunt protoconid, a well-devel-oped basal cingulid, and a crescent-shaped crest.

The p3 (Figure 3.4) is similar to the p2, but it is more cordate, wider and has a stouter aspect than p2 in occlusal view.

The p4 (Figure 3.5) is elongated with a rounded anterior mar-gin and a straight posterior edge. It is a double-rooted tooth with only one discernible cusp in antero-medial position. In the ante-rior part of protoconid there is an irregular crest. The posterior slope of the protoconid is characteristically wrinkled. There is a well-developed cingulid covering the entire base of the tooth, which becomes thinner at the most anterior edge of the tooth. There is a small thickening of the cingulid at the postero-lingual side of the tooth. The wear affects the main cusp creating a flat horizontal surface.

In the m1 (Figure 3.6) the paraconid is small. There is a short paracristid connecting the paraconid to the anterior base of the protoconid. In two specimens there is a small cingulid connect-ing the bases of the paraconid and the metaconid. Protoconid and metaconid are close to each other, they are similar in size, and they are connected by a short and robust protocristid. The talonid is larger and wider than the trigonid. The hypoconid is lower than the entoconid. The hypoconid is anteriorly con-nected to the posterior side of the protoconid by means of the oblique cristid, which is roughly parallel to the labial margin of the tooth. The entoconid is a stunt cusp and it does not connect with the metaconid. The talonid basin is open by its lingual side. The labial cingulid runs uninterruptedly from the paraconid to the posterior margin, where it raises obliquely from the base of

Figure 3. lower teeth in occlusal view of Plesiodimylus ilercavonicus sp. nov. from MaB5. 1- left c (MgUV-26747; Paratype); 2- right p1 (reversed, MgUV-26742; Paratype); 3- right p2 (reversed, MgUV-24950); 4- left p3 (MgUV-26751; Paratype); 5- left p4 (MgUV-25246; Paratype); 6- left m1 (MgUV-26737; Paratype); 7- left m2 (MgUV- 25240; Paratype). the squared area of the m2 (trigonid) is magnified at the right margin.

HISTORICAL BIOLOGY 7

concur with the latter authors that a revision of the taxonomic value of many specific characters in Plesiodimylus is needed, pay-ing special attention to the variation found in large assemblages.

The wide geographical range rapidly acquired by the genus around the MN 3–MN 4 transition is indicative of some kind of ecological advantage of Plesiodimylus over its possible com-petitors, Metacordylodon and Chainodus. Not by chance, such prosperous moment for the genus coincided with a drop in diver-sity within the rest of its family (Klietmann et al. 2014; Van den Hoek Ostende & Fejfar 2015), after the extinction of Dimylus, Dimyloides, Turkodimylus and Lacrimodon. In the light of the high diversity of morphologies found in MN 4 localities, it is deduced that during the Early Miocene the shape of the M1 in Plesiodimylus was not a character constrained by natural selec-tion. The wear pattern of several dental elements from MAB5 supports this hypothesis. The sample from this locality shows that the individuals easily wear down their occlusal reliefs with mastication, thus creating many flat surfaces, not only in M1 but also in the rest of the teeth (Figures 4 and 5). This flattening process related to the mastication involved a motion of the jaws parallel to the occlusal surface, likely a combination of protrac-tion and transverse movements of the mandible, as deduced by the presence of oblique scratches and grooves on the flat sur-faces generated (Figure 6). Thus, other than in the very early ontogenetic stages of the individuals, the minor details of the morphology of the occlusal surface were apparently not deter-minant for a successful exploitation of the feeding resources. Amblyodonty and/or hypsodonty were more likely determi-nant features involved in this issue. In that case, the taxonomy

morphological and chronological data, P. huerzeleri is the old-est and most primitive form within the genus (Klietmann et al. 2014). The genus experienced a diversification episode close to the MN 3–MN 4 transition, but it is unclear if it was a real spe-ciation process or just an intraspecific increase of the morpho-logical variability from an initial stock of P. huerzeleri. The three species P. huerzeleri, P. chantrei and P. bavaricus (if truly different species) were closely phylogenetically related. Engesser (1976) even considered P. huerzeleri to be a junior subjective synonym of P. chantrei, and many other authors followed this view (for details see Klietmann et al. 2014). On the other hand, Schötz (1985) not only considered them as different species, but he also described a new one, P. bavaricus, combining characters of both. According to Van den Hoek Ostende (1995), this latter one was not a well-defined species because it did not show any signifi-cant difference with either P. huerzeleri or P. chantrei. However, Klietmann et al. (2014) implicitly consider P. huerzeleri to be the ancestor of P. chantrei and P. bavaricus, and state advisable some kind of evolutionary connection between P. bavaricus and P. helveticus, due to the continuous presence of a mesostyle in the M1. The latter form, P. helveticus, is probably the most similar form to P. ilercavonicus nov. sp., the species described in the present paper. However, the combination of characters displayed by P. ilercavonicus sp. nov. is unknown in any other fossil sample of Plesiodimylus up to date, thus deserving the designation of a new species.

There is no doubt that Plesiodimylus encompassed an excep-tional morphological variability in its early evolutionary stages (Klietmann et al. 2014). In the large collection of Plesiodimylus from the Lower Miocene site of Petersbuch 28 (MN 3/4), with hundreds of specimens available, three morphotypes of M1 were identified, each one of them reminiscent of previously described species (namely, P. huerzeleri, P. bavaricus and P. chantrei). The authors concluded that all the elements from Petersbuch 28 were, nonetheless, better ascribed to only one species, P. aff. chantrei, a transitional evolutionary stage between P. huerzeleri and P. chant-rei. The observed variability in the upper cheek teeth was there-fore most likely explained as the evidence of an ongoing process of diversification. The slightly younger locality of Petersbuch 2 and the somewhat older ones of Erkersthofen 1 and 2 have also provided elements of Plesiodimylus ascribed to a similar transi-tional form (P. aff. chantrei).

The present case of MAB5 is certainly different to those of the Lower Miocene German localities of Petersbuch and Erkersthofen previously quoted. Even if some degree of variation is found in the morphology of the M1 (Table 2), exactly as noticed by Klietmann et al. (2014), the larger size on average, the stout aspect of all the dental elements, the complex metacone of the M2 and the ascending posterior cingulid of the m1, stand com-pletely out of the variability of the Early Miocene Plesiodimylus forms. In MAB5 there is no evidence of sediment mixing, and the range in morphology of the M1 in the assemblage is confidently explained as simple intraspecific variation. Thus, it is not unlikely that the previously known species could be indeed representing a subset of a much variable species insufficiently sampled, as it had been already proposed by Klietmann et al. (2014). On the other hand, P. chantrei may have served partly as a wastebasket taxon in the past (Doukas & Van den Hoek Ostende 2006). We

Table 2.  Variability of the characters pointed out by Klietmann et al. (2014) to discriminate M1 morphotypes observed on the material from MaB5.

M1Paras-

tyleMeso-style

Meta-style Valleys

Anterior arm of

the pro-tocone

Cingu-lum

MgUV-23385

– No rounded – – –

MgUV-23387

cusp small – 3 No con-tact

straight

MgUV-25173

– small – 3 No con-tact

curved

MgUV-25174

cusp large rounded 3 No con-tact

–

MgUV-25175

cusp small – 3 contact with paras-tyle

straight

MgUV-25182

cusp No – – – –

MgUV-25243

– small – 2 contact with paras-tyle

straight

MgUV-26734

cusp small acute 2 contact with paras-tyle

curved

MgUV-26745

cusp – – – – –

MgUV-24940

cusp – – – – –

MgUV-25208

cusp – – – – –

MgUV-25238

– – – – – straight

8 V. D. CRESPO ET AL.

Palaeoecology

The family Dimylidae has been usually regarded as indicator of humid environments because the presumed durophagous diet of its members likely restricted the distribution of these species to places where gastropods could proliferate more easily (Furió et al. 2011a; and references therein). Some authors, however, consider the genus Plesiodimylus to be a generalistic form leading

of Plesiodimylus from Central European localities could be obscured because the morphology of the M1 has been classically the reference for the identification of the species. In the model herein presented, the assemblages settled in marginal areas (like the one described in the present paper) could develop their own morphological features more easily simply because they were more susceptible of getting isolated from the source area, thus precluding the genetic flow.

Figure 4. Upper teeth in labial view of Plesiodimylus ilercavonicus sp. nov. from MaB5, showing different states of wear. the upper row (1–4) includes teeth of young individuals with no wear or in an early stage of wearing (1- right c reversed, MgUV-27027; 2- right P4 reversed, MgUV-25244; 3- right M1 reversed, MgUV-23387; 4- left M2, MgUV-26735). the intermediate row (5–8) includes teeth of adults with a moderate wear of their occlusal surfaces (5- left c, MgUV-25241; 6- right P4 reversed, MgUV-25195; 7- left M1, MgUV-26734; 8- left M2, MgUV-26637). the lower row (9–10) show two teeth of old-aged individuals with a severe degree of wear (9- right c reversed, MgUV-25204; 10- right M1 reversed, MgUV-25173). in them, their occlusal morphologies are hardly identifiable because the wear has created large flat surfaces.

Figure 5. lower teeth in labial view of Plesiodimylus ilercavonicus sp. nov. from MaB5, showing different states of wear. the upper row (1–5) includes teeth of young individuals with no wear or in an early stage of wearing (1-left c, MgUV-26747-Paratype; 2-right p1 reversed, MgUV-26742-Paratype; 3-left p4, MgUV-25246-Paratype; 4-left m1, MgUV-25159; 5-left m2, MgUV-25240-Paratype). the intermediate row (6–9) includes teeth of adults with a moderate wear of their occlusal surfaces (6-right c reversed, MgUV-26755; 7-right p4 reversed, MgUV-26741; 8-left m1, MgUV-26737; 9-left m2, MgUV-26738). the lower row (10–13) shows four teeth of old-aged individuals with extreme degrees of wear (10-left c, MgUV-26748; 11-right p1 reversed, MgUV-26743; 12-right p4 reversed, MgUV-26998; 13-right m1 reversed, MgUV-26739). the flat surfaces on their occlusal sides are remarkable.

HISTORICAL BIOLOGY 9

During all its temporal range, the presence of Plesiodimylus is more common and continuous in Central Europe (where it is to be considered a resilient form) than in the southern areas (where it is rather a transient genus) (Van den Hoek Ostende et al. 2016). In the inner zones from Europe, the genus has been reported from many MN 3 to MN 10 localities from France (e.g., Baudelot 1972; Mein 1999; Mein & Gingsburg 2002; Engesser 2009), Germany (e.g., Müller 1967; Ziegler & Fahlbusch 1986; Ziegler 1995; Ziegler & Mörs 2000; Ziegler et al. 2005; Seehuber 2008; Prieto 2011), Poland (e.g., Rzebik-Kowalska 1996, 2009), Ukraine (e.g., Nesin & Topachevsky 1999; Nesin & Nadachowski 2001), Switzerland (e.g., Hürzeler 1944; Engesser 1972; Kälin & Engesser 2001), Hungary (e.g., Prieto et al. 2012; Hír et al. 2016), Austria (e.g., Bachmayer & Wilson 1990; Ziegler & Daxner-Höck 2005; Ziegler 2006), Czechia and Slovakia (e.g., Fejfar & Sabol 2005). In contrast, the occurrences in southern Europe have been only documented during the Early Miocene (MN 4) and the late Middle Miocene (MN 7  +  8). During the Early Miocene (MN 4) the genus reached Greece (Doukas 1986; Doukas & Van den Hoek Ostende 2006), where P. chantrei and P. aff. crassidens are found, and Spain (present work). During the late Middle Miocene (MN 7 + 8), the genus reached Turkey and Spain. In Turkey only P. crassidens has been reported (Engesser 1980). In Spain, P. chantrei is apparently the only species present in the localities from the Vallès-Penedès Basin, ranging from MN 7+8 to MN 10 from Terrassa (Van den Hoek Ostende & Furió 2005; Furió et al. 2011a, 2011b).

The distribution of this genus was much likely linked to the Miocene climatic fluctuations affecting both precipitation and temperatures. The end of the Early Miocene (MN4, upper Burdigalian) was a moment of frequent rainfalls and high

an insectivorous diet as those of erinaceids or some talpids, there-fore being the less specialized genus of this family (Ziegler 2005). Actually, this statement seems valid for those species with rather faint teeth (P. hurzeleri, P. chantrei, and P. bavaricus), whereas the forms with more amblyodont dentitions included in the genus (P. similis, P. crassidens, P. helveticus, P. johanni, P. gaillardi, and P. ilercavonicus sp. nov.) followed the general rule for the family.

The dental microwear is indicative of the prevailing diet in several mammalian orders (Ungar 2015). Unfortunately, most of the studies have been focused on primates, ruminants and rodents, being the diets of insectivores (i.e., eulipotyphlans) and their corresponding wear patterns poorly known so far. To our knowledge, only the dental microwear of some talpids (Silcox & Teaford 2002) and soricids (Withnell & Ungar 2014) has been preliminary studied to discriminate the patterns left by earth-worms and vertebrates over mostly insect-feeders. The general rule for mammals states that tough-food eaters have microwear surfaces dominated by uniform scratches running in a given direction and hard-object feeders show pits of different sizes and shapes (Ungar 2015). Both statements are apparently valid when they are applicated at different scales to the teeth of P. ilercavon-icus nov. sp. Whereas at a magnification to 500 μm sub-parallel scratches are dominant, a higher magnification till the 100 μm make pits look preferential (Figure 6). This approach is therefore not conclusive, but worthy to be further developed in a future.

The evolution of the genus in time and space can provide some extra clues on the most prolific environments for Plesiodimylus. The oldest known occurrences of the genus are reported from the MN3 localities of Estrepouy (France; Hugueney & Bulot 2011), Wintershof-West (Germany; Müller 1967) and Ahníkov 1 (Czech Republic; Van den Hoek Ostende & Fejfar 2015).

Figure 6. Detailed sEM photograph of the wear flat surface generated on the occlusal side of a right p4 (MgUV–26998). right: Notice that the scratches are obliquely oriented, sometimes almost transverse to the mandibular axis, but they never get a complete longitudinal direction. Middle: some pits are discernible in the sEM photograph at 100 μm. left: some minor scratches are visible with magnification at 10 μm.

10 V. D. CRESPO ET AL.

FundingThis work was financially supported by the support of the Ministerio de Economía y Competitividad (Government of Spain, Projects CGL2015-63777-P and CGL2016-76431-P), and the help of Generalitat de Catalunya by means of CERCA Program and projects 2014 SGR 416 GRC (Agència de Gestió d’Ajuts Universitaris i de Recerca, AGAUR) and 2014/100584-‘Canvis Climàtic i Faunístics al Miocè Inferior de Catalunya’ (Departament de Cultura). The campaigns of prospection and excava-tion in the area of Araia d’Alcora have been funded by the Conselleria de Cultura y Esports of the Valencian government since 2008–2011.

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temperatures (Böhme 2003) that preceded the strong season-ality (and dryness in Central Iberia) of the Miocene Climatic Optimum (upper Burdigalian–Langhian) (Mosbrugger et al. 2005; Jiménez-Moreno & Suc 2007). Purportedly, the high temperatures and precipitation triggered the northern migra-tion of termophilic ectothermic vertebrates and the presence of paratropical rainforest (Böhme 2003). This fact agrees with the presence of the genus Plesiodimylus in the south of Europe during the MN 4 and its absence in most of the MN 5 and MN 6 Spanish localities (Van den Hoek Ostende et al. 2016). The second expansion to the south of Europe during the late Middle Miocene (MN 7 + 8) also coincides with another humid period in Europe during the late Serravallian-Tortonian (Bruch et al. 2011).

The family Dimylidae went finally extinct in the transition from the Vallesian to the Turolian (MN 10 – MN 11), after a long period of decline, likely due to the aridification, the high tem-peratures, the increase of the open vegetation of the Tortonian (Bruch et al. 2011). The last documented occurrences correspond to MN 11 localities of Ambérieu 3 (Mein 1999), and Dorn-Dürkheim 1 (Ziegler 2005) from Central Europe. There is not yet a satisfactory explanation for its final extinction, though it is a well known fact that its ecological niche was rapidly occupied by the soricid Amblycoptus (a putative descendant genus of a ‘Crusafontina stock’, according to Van Dam 2010), which emu-lated most of the differential traits of Plesiodimylus.

Conclusions

The Plesiodimylus assemblage from Mas d’Antolino B 5 repre-sents the first record of this genus from Lower Miocene on the Iberian Peninsula. The amblyodont teeth indicates that P. iler-cavonicus sp. nov. was one of the most malacophagous species within the genus. Its presence in the Iberian Peninsula during the Early Miocene coincides with the expansion of favourable environmental conditions like those present in Central Europe during most of the Miocene. The isolation of this population in a marginal area probably enhanced the speciation processes and the development of differential traits respect to the Central European dwellers.

AcknowledgmentsMF acknowledges the financial support of the Ministerio de Economía y Competitividad (Government of Spain, Projects CGL2015-63777-P and CGL2016-76431-P), and the help of Generalitat de Catalunya by means of CERCA Program and projects 2014 SGR 416 GRC (Agència de Gestió d’Ajuts Universitaris i de Recerca, AGAUR) and 2014/100584-‘Canvis Climàtic i Faunístics al Miocè Inferior de Catalunya’ (Departament de Cultura). The campaigns of prospection and excavation in the area of Araia d’Alcora have been funded by the Conselleria de Cultura y Esports of the Valencian government since 2008–2011. We appreciate the help of C. Doukas, M. Freudenthal, R. Ziegler, and S. Mansino, who provided literature of difficult access, and O. Suárez-Hernando and X. Murelaga, who performed the stra-tigraphy of the MAB section. We kindly appreciate the technical assistance provided by the staff members from the Microscopy Section of the SCSIE (UV) and the helpful comments on the original manuscript provided by editor Gareth Dyke, R. Ziegler and an anonymous reviewer.

Disclosure statementNo potential conflict of interest was reported by the authors.

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AbstractIntroductionGeographic and Geologic SettingMaterial, methods and abbreviationsSystematic paleontologyDifferential DiagnosisSynonymyMaxillaryUpper DentitionMandibleLower DentitionTeeth of difficult identification

Description of the materialDiscussionPalaeoecologyConclusionsAcknowledgmentsDisclosure statementFundingReferences