Desmaninae (Talpidae, Mammalia) from the Pliocene of Tollo de Chiclana (Guadix Basin, Southern Spain)

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Geobios 41 (2008) 381–398 http://france.elsevier.com/direct/GEOBIO/

Original article

Desmaninae (Talpidae, Mammalia) from the Pliocene of Tollo de Chiclana (Guadix Basin, Southern Spain) Considerations on the phylogeny of the genus Archaeodesmana Desmaninae (Talpidae, Mammalia) du Plioce`ne de Tollo de Chiclana (Bassin de Guadix, Espagne me´ridionale) Conside´rations sur la phyloge´nie du genre Archaeodesmana Raef Minwer-Barakat a,*, Antonio García-Alix a, Matthijs Freudenthal a,b a

Departamento de Estratigrafía y Paleontología, Universidad de Granada, Facultad de Ciencias, Avda. Fuentenueva s/n, 18071 Granada, Spain b Nationaal Natuurhistorisch Museum, Naturalis, P.O. Box 9517, 2300 RA Leiden, The Netherlands Received 23 May 2007; accepted 9 August 2007 Available online 4 March 2008

Abstract The fossil Desmaninae (water-moles) from the Pliocene continental deposits of Tollo de Chiclana (Guadix Basin, Southern Spain) are described. A new species, Archaeodesmana elvirae, is defined from the locality of Tollo de Chiclana-1 (upper Ruscinian). This species is characterized by relatively small canines and premolars (except the P4) and large P4 and molars, besides several morphological features. The presence of Archaeodesmana brailloni is reported from the locality of Tollo de Chiclana-1B (uppermost Ruscinian). A small sample assigned to the genus Archaeodesmana is described from the lower Villafranchian site of Tollo de Chiclana-3, which cannot be determined at the specific level. The phylogenetic relationships between the different species of Archaeodesmana are reconsidered in the light of the recent findings, which support the idea of a more complex phylogeny than previously proposed for this genus. The populations from the Guadix Basin, previously assigned to Dibolia dekkersi (= Archaeodesmana getica), are here considered to belong to a different (unnamed) species, which is the ancestor of A. elvirae. On the other hand, the new species A. elvirae is proposed as the ancestor of A. brailloni. # 2008 Elsevier Masson SAS. All rights reserved. Résumé Parmi les Desmaninae fossiles du Pliocène continental de Tollo de Chiclana (Bassin de Guadix, Espagne méridionale) une nouvelle espèce, Archaeodesmana elvirae, est reconnue dans le gisement de Tollo de Chiclana-1 (Ruscinian Supérieur) ; cette espèce est caractérisée par des canines et des prémolaires (excepté la P4) relativement petites, par une P4 et des molaires de grande taille, et par quelques caractères morphologiques. Archaeodesmana brailloni est documentée au gisement Tollo-de-Chiclana-1B (Ruscinian final). Le site du villafranchien inférieur, Tollo-deChiclana-3, a livré une petite collection de Archaeodesmana, qui ne peut pas être déterminée au niveau spécifique. Les relations phylogénétiques entre les espèces de Archaeodesmana sont révisées, ce qui mène à une conception plus complexe que supposée jusqu’à présent. Les populations du Bassin-de-Guadix, attribuées auparavant à Dibolia dekkersi (= Archaeodesmana getica) sont considérées comme représentant une autre espèce — non nommée — qui est l’ancêtre de A. elvirae. Elle est également proposée comme ancêtre de A. brailloni. # 2008 Elsevier Masson SAS. All rights reserved. Keywords: Talpidae; Desmaninae; Archaeodesmana; Neogene; Ruscinian; Spain Mots clés : Talpidae ; Desmaninae ; Archaeodesmana ; Néogène ; Ruscinien ; Espagne

* Corresponding author. E-mail address: [email protected] (R. Minwer-Barakat). 0016-6995/$ – see front matter # 2008 Elsevier Masson SAS. All rights reserved. doi:10.1016/j.geobios.2007.08.001

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1. Introduction The subfamily Desmaninae (desmans) includes only two extant species: Galemys pyrenaicus and Desmana moschata, both with a semi-aquatic lifestyle. G. pyrenaicus inhabits cold mountain lakes and rivers in the northern part of the Iberian Peninsula and the Pyrenees mountains. D. moschata prefers slow moving streams, lakes and ponds; this species occupies the Volga, Ural, Don and Oka Rivers, in Eastern Europe. Water-moles show anatomic adaptations for swimming and diving: the hind legs are much longer than the front legs, the feet are webbed, etc. Some of these features can also been observed in fossil species (Rümke, 1985a; Martín Suárez et al., 2001). Extant water-moles are always related with permanent streams. Inferring the same requirements for fossil Desmaninae, it is clear that this mammal group has a great paleoecological relevance, since its presence can be associated with relatively humid and rainy conditions. Several Miocene and Pliocene water-mole species were grouped by Rümke (1985a) under the name of Dibolia. Posteriorly, Rzebik-Kowalska and Pawlowski (1994) pointed out that this name is a junior homonym of Dibolia Latreille, 1829, a genus of Coleoptera and proposed the generic name Ruemkelia. Later, Hutterer (1995) elevated the subgenus Archaeodesmana Topachevski and Pashkov, 1983 to generic rank and included nine species:         

Desmana pontica Schreuder, 1940 (type species) Desmana vinea Storch, 1978 Desmana bifida Engesser, 1980 Dibolia dekkersi Rümke, 1985a Dibolia turolense Rümke, 1985a Dibolia adroveri Rümke, 1985a Dibolia brailloni Rümke, 1985a Dibolia luteyni Rümke, 1985a Dibolia major Rümke, 1985a.

We accept the nomenclatural revision of Hutterer (1995), so Ruemkelia is considered a junior synonym of Archaeodesmana. Martín Suárez et al. (2001) included Desmana verestchagini Topachevski, 1961 in this genus and a new species, Archaeodesmana baetica. Desmana getica Terzea, 1980 was considered a junior synonym of A. bifida by Rümke (1985a), but was transferred to A. dekkersi by Van den Hoek Ostende et al. (1989). Provided these authors are right, dekkersi would be a junior synonym of getica. Following Rzebik-Kowalska (2005), we consider that this species is also included in the genus Archaeodesmana. The diagnostic characters of the genus Archaeodesmana are: I1 is short and straight with a bifid tip, in I1, the crown is large and does not overhang the root anteriorly; P2 is usually longer than P3; P2 is always longer than P3 (Rümke, 1985a). Despite its abundance, the genus Archaeodesmana has been poorly studied in the Iberian fossil record: there are very few papers dealing with this genus and other fossil Desmaninae and their representatives are often not identified at specific level. In

this paper we describe three populations ascribed to Archaeodesmana, from the localities of Tollo de Chiclana-1, 1B and 3 (TCH-1, 1B and 3), situated in the Guadix Basin (Southern Spain). We describe a new species, Archaeodesmana elvirae, from TCH-1 and we reconsider the phylogenetic relationships between the different species of this genus proposed in previous papers. 2. Geological setting The Guadix Basin is situated in the Betic Cordillera (Spain), one of the mountain chains formed during the closing of the Tethys (Sanz de Galdeano, 1990). It was established as a separate intramontane basin in the late Miocene (Viseras et al., 2004, 2005). The oldest sedimentary infilling of this basin was deposited in a phase of marine sedimentation during the Tortonian (Fernández et al., 1996; Soria et al., 1999). The upper part of the infilling represents a stage of exclusively continental sedimentation that lasted from latest Tortonian until the Late Pleistocene (Viseras, 1991). The area of Tollo de Chiclana is situated in the central part of the basin, where fine-grained detritic deposits, corresponding to distal fluvial sedimentation, alternate with lacustrine carbonates (Viseras, 1991; Fernández et al., 1996). From this area, several superposed Pliocene fossiliferous localities were described in previous publications (Minwer-Barakat et al., 2004, 2005). These fossiliferous levels are found in swamp deposits and have yielded rich micromammal faunas composed of various families of rodents and insectivores (Minwer-Barakat, 2005). Remains ascribed to the genus Archaeodesmana have been found in the localities of Tollo de Chiclana-1, Tollo de Chiclana-1B (late Ruscinian) and Tollo de Chiclana-3 (early Villafranchian). In younger localities (Tollo de Chiclana-13, 10 and 10B), remains of Desmaninae have been found that cannot be determined at generic level; the latter material will not be described in the present paper. 3. Dental terminology and measuring methods The nomenclature used in the descriptions of the teeth is that of Rümke (1985a). Lengths and widths have been measured as defined by Martín Suárez et al. (2001). Measurements were only taken on canines, premolars and molars because data on incisors are rarely reported in the literature. In the lower molars, only the length was measured, because the width is greatly influenced by orientation and seems not to be a reliable parameter for making comparisons. The specimens are kept in the ‘Departamento de Estratigrafía y Paleontología’ of the University of Granada, Spain. 4. Systematic Paleontology Family TALPIDAE Fischer von Waldheim, 1817 Subfamily DESMANINAE Mivart, 1871 Genus Archaeodesmana Topachevski and Pashkov, 1983 Type species: D. pontica Schreuder, 1940.

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Archaeodesmana elvirae nov. sp. Fig. 1 Etymology: Dedicated to Dr. Elvira Martín Suárez (University of Granada), who discovered the fossiliferous localities of Tollo de Chiclana, in recognition of her outstanding work in vertebrate paleontology. Type locality: Tollo de Chiclana-1 (TCH-1), late Ruscinian, Guadix Basin, Spain. Coordinates UTM 30SVG959553 (Minwer-Barakat, 2005). Holotype: Left M1, TCH-1 212, deposited in the ‘Departamento de Estratigrafía y Paleontología’, University of Granada. Referred specimens: 4 I1 (TCH-1 158-161); 5 I2 (TCH1 162–166); 3 P1 (TCH-1 167–169); 1 P2 (TCH-1 170); 3 P3 (TCH-1 171–173); 3 P4 (TCH-1 174–176); 7 M1 (TCH-1 177– 183); 9 M2 (TCH-1 184–192); 8 M3 (TCH-1 193–200); 2 I1 (TCH-1 240; TCH-1 270); 1 I2–3 (TCH-1 201); 1 C sup. (TCH1 202); 1 P1 (TCH-1 203); 2 P2 (TCH-1 204, 205); 5 P4 (TCH1 206–210); 7 M1 (TCH-1 211–217); 10 M2 (TCH-1 218–227); 12 M3 (TCH-1 228–239). Diagnosis: Archaeodesmana with relatively small premolars (except P4 and P4), similar in size to those of A. getica, and with large P4 and molars, similar in size to those of A. brailloni. Lower premolars with weak cingulids. P1 with two fused roots. P4 with a distinct paraconid. Lower molars with a long and wide anterolabial cingulid and with a prominent parastylid. M1 and M2 with a protruding entostylid. P2 long, without lingual expansion. P4 strongly molarized, with large, prominent protocone, wide parastyle and incipient metacone. Upper molars with a well-individualized accessory cusp. M1 with prominent parastyle and metastyle. Differential diagnosis: A. elvirae differs from A. getica in its larger size, especially in the P4 and the molars, in the narrower posterior cingulid in the lower molars and premolars and in the two fused roots in the P1. The new species differs from A. vinea in its notably larger size, mainly in P2, P4 and the molars and in the lower ratio between the mean length of P2 and P3; in addition, A. elvirae differs from A. vinea in the P2 with two roots and without a distinct lingual extension. Biometrically, A. elvirae can be distinguished from A. turolensis by its smaller P2 and P2 and by its larger M1, M2, P4 and M1. Archaeodesmana adroveri has clearly larger P2, P3 and P4 and a shorter upper canine than A. elvirae. All the premolars of A. major, except the P4, are markedly larger than those of A. elvirae, whereas the P4 and the M1 are smaller. Morphologically, the lower molars of A. turolensis, A. adroveri and A. major have less-developed parastylids than those of A. elvirae and the upper molars have a smaller protoconule and a minuscule or even absent accessory cusp. Archaeodesmana pontica has notably larger premolars and a smaller M1 than A. elvirae; in addition, the P2 of A. pontica has a lingual extension and three roots. Archaeodesmana bifida differs from A. elvirae in the P2 shorter than the P3; besides, its P1, P4, M1 and M2 are smaller than those of A. elvirae, its lower molars lack a parastylid, its I1 is narrower and its P4 has a lessdeveloped protocone and lacks a metacone. The P2, P4 and the upper canine of A. elvirae are much smaller than those of

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A. baetica; moreover, the premolars of A. baetica are less molarized, the lower premolars have less-developed cingulids and the M3 have less-developed lingual cusps than those of A. elvirae. A. elvirae differs from A. brailloni in the notably smaller P2, P3 and P2, in the less-molarized lower premolars, in the more-developed parastylid in the lower molars and in the lack of a lingual extension in the P2. The molars and the P4 of A. luteyni are smaller than those of A. elvirae and the P4 lacks a paraconid and metaconid. A. elvirae can be distinguished from A. verestchagini by its much smaller premolars, except for P4; the P2 of A. verestchagini has an accessory lingual root and the upper molars have smaller protoconules and metaconules than those of A. elvirae. Occurrence in the studied area: Tollo de Chiclana-1 (TCH-1), late Ruscinian. Measurements: See Table 1. Description: I1: The crown is shovel-shaped; its base has a triangular cross-section. The enamel extends just as far down anteriorly as posteriorly (morphotype ‘‘a’’ of Rümke, 1985a). There is a pronounced posterior projection, bordered by a narrow cingulid. A rib crosses the posterior face vertically. The root is long and straight, with triangular cross-section. I2: The crown is higher than in I1, compressed at the apex. The labial face is slightly convex; the lingual one is convex in its upper part and concave near the base. The posterocristid is sharp and long. There is a narrow but well-differentiated cingulid bordering the lingual and posterior faces. At the intersection between the cingulid and the posterocristid, there is a small, hook-shaped cuspid. The root is very long, with rounded cross-section. P1: The crown is low. The labial border is rounded; the lingual, anterior and posterior borders are pointed. The labial face is convex; the lingual one is flat and, towards its base, it becomes slightly concave. The cingulid is weak and occupies the lingual border of the tooth, widening in its central part. The anterocristid is sharp in two specimens and blunt in the third one. There is no posterocristid. There are two fused roots. P2: The crown is notably higher than in P1. The labial and lingual faces are convex. The anterolabial part of the tooth is flat; the anterolingual extreme is pointed. The anterocristid is well-marked, extends to the anterolingual end of the tooth and reaches a small anterolingual cingulid. The posterocristid is limited to the highest part of the protoconid. The posterior cingulid is weak and narrow, with a small bulge at the posterolingual end of the tooth. There is no lingual cingulid. The tooth is poorly molarized: there is neither a talonid basin, nor a distinct paraconid. The roots are not preserved. P3: The crown is lower than in P2. The anterolabial, anterolingual and posterolingual borders are straight and the posterolabial one is curved. The anterocristid is sharp and reaches the base of the crown. The posterocristid is limited to the upper part of the protoconid. In two out of three specimens, there is a weak enamel bulge at the anterior end of the tooth. The cingulid is wide in the posterior part of the tooth and becomes narrower towards the labial face. There is no lingual

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Fig. 1. A. elvirae nov. sp. from TCH-1. A, right I1, TCH-1 160. B, right I1, TCH-1 240. C, right I2, TCH-1 164. D, right P1, TCH-1 169. E, right P2, TCH-1 170. F, right P3, TCH-1 173. G, right P4, TCH-1 176. H, left I2–3, TCH-1 201. I, left C sup., TCH-1 202. J, right P2, TCH-1 204. K, left P4, TCH-1 206. L, right M3, TCH1 198. M, right M2, TCH-1 190. N, right M1, TCH-1 178. O, left M1, TCH-1 212 (holotype). P, left M2, TCH-1 221. Q, left M3, TCH-1 228. A–C, posterior view. D–Q, occlusal view. Scale bar = 1 mm. Fig. 1. A. elvirae nov. sp. de TCH-1. A, I1 droite, TCH-1 160. B, I1 droite, TCH-1 240. C, I2 droite, TCH-1 164. D, P1 droite, TCH-1 169. E, P2 droite, TCH-1 170. F, P3 droite, TCH-1 173. G, P4 droite, TCH-1 176. H, I2–3 gauche, TCH-1 201. I, C sup. gauche, TCH-1 202. J, P2 droite, TCH-1 204. K, P4 gauche, TCH-1 206. L, M3 droite, TCH-1 198. M, M2 droite, TCH-1 190. N, M1 droite, TCH-1 178. O, M1 gauche, TCH-1 212 (holotype). P, M2 gauche, TCH-1 221. Q, M3 gauche, TCH-1 228. A–C, vue postérieure. D–Q, vue occlusale. Échelle = 1 mm.

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Table 1 Measurements (in mm) of A. elvirae nov. sp. from TCH-1, A. brailloni Rümke, 1985a from TCH-1B and Archaeodesmana sp. from TCH-3 Tableau 1 Mesures (en mm) de A. elvirae nov. sp. de TCH-1, A. brailloni Rümke, 1985a de TCH-1B et Archaeodesmana sp. de TCH-3 Length

C inf. P1 P2 P3

P4 M1 M2 M3

C sup. P1 P2 P3 P4

M1 M2 M3

TCH-1B TCH-1B TCH-1 TCH-1B TCH-1 TCH-3 TCH-1B TCH-1 TCH-1B TCH-1 TCH-1B TCH-1 TCH-1B TCH-1 TCH-3 TCH-1B TCH-1 TCH-1B TCH-1 TCH-1 TCH-1B TCH-1 TCH-3 TCH-1B TCH-3 TCH-1B TCH-1 TCH-1B TCH-1 TCH-1B TCH-1 TCH-1B TCH-1

Width

n

Minimum

Mean

Maximum

n

Minimum

Mean

Maximum

4 4 3 3 1 1 4 3 4 2 4 1 5 6 1 7 7 3 1 1 2 2 1 2 1 1 2 1 4 4 6 4 6

0.96 1.12 0.93 1.49

1.00 1.18 0.97 1.55 1.37 1.29 1.48 1.26 1.64 1.665 2.58 2.61 2.50 2.48 1.91 2.03 1.95 1.38 1.34 1.17 1.835 1.645 1.34 1.56 2.28 2.20 2.31 3.30 3.28 2.27 2.25 1.59 1.47

1.07 1.23 1.01 1.59

4 4 3 3 1 1 4 3 4 2

0.86 0.99 0.82 1.06

0.90 0.995 0.86 1.08 0.94 0.85 1.05 0.90 1.20 1.065

0.93 1.01 0.88 1.10

3 1 1 2 2 1 3 1 1 1 1 3 4 6 4 9

1.09

1.10 0.93 0.88 1.23 1.01 1.17 1.28 1.80 1.92 1.87 2.49 2.36 2.72 2.655 2.185 2.07

1.11

1.45 1.20 1.61 1.63 2.54 2.45 2.44 1.98 1.90 1.33

1.83 1.60 1.54

2.30 3.14 2.21 2.16 1.53 1.39

cingulid. The available specimens only preserve the anterior root, which has a round cross-section. P4: Long and narrow teeth, with a very high crown. The lingual and posterior borders are straight; the labial and anterior borders are curved. The anterocristid is long and sharp; it is connected to a small, well-individualized paraconid. There is an incipient metaconid. The posterior part of the tooth is variable: in TCH-1 176, the posterocristid occupies only the upper half of the crown; there is another crest in a more lingual position, which connects to a well-differentiated entoconid. In TCH-1 175, the posterocristid reaches the posterior border; this specimen has another, shorter, crest in a more lingual position, which does not reach the base of the crown and lacks an entoconid. The specimen TCH-1 174 has a distinct entoconid, but lacks a posterocristid. All the specimens have a robust cingulid, limited to the posterior border. There is also a narrower anterior cingulid. The talonid basin is very small and lingually open. There are two roots. M1: The anterolabial cingulid is wide, especially in its anterior portion; it reaches the base of the hypoconid without interruptions. Two out of three specimens preserving the anterior part have a well-developed parastylid; in the third one,

1.53 1.31 1.68 1.70 2.62

1.03 0.86 1.14 1.05

1.06 0.93 1.25 1.08

2.57 2.51 2.14 2.01 1.43

1.84 1.69 1.58

2.32 3.40 2.35 2.36 1.65 1.59

1.21 1.00 1.25

2.28 2.65 2.57 2.12 2.01

1.25 1.02 1.30

2.41 2.80 2.71 2.23 2.18

the cingulid is widened at the anterolingual end. The oblique cristid reaches the top of the metaconid. The posterior cingulid is short and narrow. The entostylid is small but prominent. None of the specimens preserves the anterior root; the posterior root has a deep groove on its internal face. M2: The anterolabial cingulid is variable in width, continuous and reaches the hypoconid. This cingulid rises notably at the level of the re-entrant valley, forming a triangular-shaped protuberance. The parastylid is large and prominent. The oblique cristid reaches the top of the metaconid. The posterior cingulid is very narrow, in some specimens almost indiscernible. The entostylid is very prominent and smaller than the parastylid. There are two roots. M3: The protoconid and metaconid are high and sharp; the hypoconid and entoconid are rounded. The anterolabial cingulid is narrow and reaches the base of the hypoconid. The parastylid is very wide and prominent. The oblique cristid reaches the top of the metaconid. There is neither a posterior cingulid nor an entostylid. The posterior border is rounded. There are two roots. I1: The crown is low, with a triangular occlusal outline. The anterior and labial faces are slightly convex and the lingual face

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is strongly concave. Due to the preservation of the specimens, neither the indentation of the anterior border nor the cingulum can be observed. The tooth is divided into two lobes, the labial one larger than the medial one. The enamel is much thicker in the anterior and labial faces than in the lingual one. There is an enamel-free area in the medial face. The medial crest is short. The root, preserved in one specimen only, is straight and strongly compressed. I2–3: The crown is small, conical, with nearly circular crosssection. The labial face is slightly convex; the lingual face is flat. The anterocrista and the posterocrista are very faint. The cingulum is narrow and poorly defined; it occupies the entire lingual border and it broadens at the anterior and posterior ends of the tooth. The root is strongly compressed transversally and fan-shaped. C sup.: The crown is high and conical. The outline is elliptical. The anterior and posterior parts are similar in width. The enamel base rises at the middle of the labial and lingual faces. There is no anterocrista. The sharp posterocrista does not reach the base of the crown. The anterior and posterior cingula are wide, especially at the anterolingual and posterolingual ends of the tooth and connected lingually. There are two divergent roots. P1: The crown is low, with elliptical cross-section. The labial face is convex; the lingual one is slightly concave. No cingula can be observed, but that may be due to the poor preservation of the base of the crown. The posterocrista is long and weak. There is no anterocrista. The roots are not preserved. P2: The crown is high and conical. The outline is elliptical; the length is notably larger than the width. The labial face is convex. The anterolingual and posterolingual parts are flattened. The anterior cingulum is narrow and restricted to the anterior border. The posterior cingulum is slightly wider and borders the posterior third part of the labial face, the posterior face and the posterior half of the lingual face. The enamel base rises at the centre of the lingual and labial faces. The posterocrista does not reach the cingulum. There are two roots. P4: Only one out of the five specimens is complete. The labial border is rounded, the posterolingual border is slightly concave and the anterolingual border is strongly concave. There is no anterocrista. The posterocrista is sharp, widening near the posterior border and forming a small metacone, which is well individualized in two specimens only. The protocone is large and distinct and forms a prominent lingual expansion; in one specimen, it is connected to the paracone by a small crest. The parastyle is very wide, prominent and connected to an anterolabial cingulum and to an anterolingual cingulum; both cingula are narrow and poorly defined. The metastyle is narrower than the parastyle. The posterolabial cingulum is high and well marked; it delimits a small basin at the posterolabial base of the paracone. The posterolingual cingulum is wide and strong. In two specimens, it ends at the base of the protocone; in the other three, it reaches the apex of the protocone. There are three roots: below the anterior, posterior and lingual ends of the tooth. M1: The parastyle is very prominent; it is connected to the paracone by a weak and low posterior crest and to the protoconule by a somewhat wider anterior cingulum. The

metastyle is narrow and elongated. The protoconule is separated from the protocone by a deep groove. The accessory cusp, attached to the posterior part of the protocone, is small and rounded. The metaconule is equal in size to the protoconule and separated from the accessory cusp. The posterior cingulum is long, narrow and continuous. There is a very short cingulum below the valley that separates the protocone from the protoconule and there is another, longer, narrow cingulum below the valley that separates the protocone from the metaconule. There are three main roots in lingual, anterolabial and posterolabial position and a small secondary root below the centre of the crown. M2: The parastyle is a mere widening of the anterolabial cingulum. The metastyle is somewhat narrower than the parastyle. The protoconule is separated from the protocone by a deep groove. At the base of the groove, there is a minuscule, but distinguishable, rounded tubercle. The protocone is large and markedly shifted towards the anterolingual part. The accessory cusp is larger than in M1. The metaconule is somewhat higher than the protoconule, separated from the metacone and from the accessory cusp by deep valleys. The anterior cingulum is narrow, continuous and well separated from the paracone. The posterior cingulum is weak, low and attached to the metacone. There are three main roots in anterolabial, posterolabial and lingual position and a small secondary root below the centre of the tooth, connected to the main roots by three crests. Contrary to M1, there is another secondary root, small and rounded, below the centre of the labial border. M3: The parastyle is absent in three out of 12 specimens; in the rest it is a slight widening of the anterior cingulum. The posterior mesostyle is poorly individualized. The protoconule is well separated from the protocone by a wide and deep groove. There is a small and low accessory cusp, attached to the posterior part of the protocone. The metaconule is well individualized, larger than the protoconule and separated from the metacone and from the accessory cusp by two deep valleys. One specimen has an additional small, rounded cusp on the posterior border, between the metacone and the metaconule. The anterior cingulum is very narrow and attached to the paracone, or even absent in three out of 10 specimens preserving the anterior part. There is a minuscule enamel protuberance or a very short cingulum at the base of the crown, between the protocone and the protoconule. There are three roots in lingual, anterolabial and posterolabial position. Comparisons: The population from TCH-1 cannot be ascribed to any of the known species of Archaeodesmana, which are distinguished mainly on the basis of biometrical criteria (Rümke, 1985a). That kind of criteria is sufficient to differentiate the population of A. elvirae from TCH-1 from practically all the species included in the genus; in some cases the differences in size are very marked. However, we have also taken into account the morphology of the teeth, especially in the cases in which the biometrical differences are subtle. The size of A. vinea (Storch, 1978), from the localities of Dorn-Dürkheim and Kohfidisch, is notably smaller than that of A. elvirae from TCH-1 for most dental elements, especially P2, P4 and molars (see Tables 1 and 2). Moreover, the ratio between

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Table 2 Mean lengths (in mm) of several teeth used for the diagnosis of the different species of Archaeodesmana Tableau 2 Longueurs moyennes (en mm) des éléments dentaires diagnostiques des différentes espèces de Archaeodesmana A. geticaa A. vinea A. turolensis A. pontica A. adroveri A. bifidab A. baetica A. brailloni A. luteyni A. major A. verestchagini A. elvirae P2 P3 P4 C sup P2 P4 M1

1.44 1.24 1.58 1.28 1.59 1.99 2.79

1.45 1.15 1.62 1.49 1.92 2.95

1.52 1.32 1.65 1.41 1.76 2.16 2.96

1.64

1.38 1.79 3.02

1.60 1.41 1.82 1.21 1.78 2.29 3.12

1.32 1.40 1.66 1.40 2.06 3.09

2.03 1.43 2.18 1.68 1.70 2.35 3.19

1.49 1.44 1.70 1.36 1.77 2.24 3.09

1.25 1.65 1.39 1.59 2.01 2.99

1.56 1.43 1.72 1.58 1.93 2.12 2.99

1.84 1.53 1.97 1.33 1.98 2.30 3.04

1.37 1.26 1.67 1.34 1.65 2.31 3.28

Data for A. baetica from Martín Suárez et al. (2001); for A. pontica and A. getica from Rümke (1985a); for A. elvirae this paper; other data from Rümke (1985b). Données pour A. baetica de Martín Suárez et al. (2001) ; pour A. pontica et A. getica de Rümke (1985a) ; pour A. elvirae ce travail ; autres données de Rümke (1985b). a The measurements of A. getica are based only on the assemblage from Kardia. b The measurements of A. bifida are based only on the assemblage from Dinar-Akçaköy.

the mean lengths of the P2 and the P3 is much higher in A. vinea than in A. elvirae. The P2 of A. vinea, with a well-marked lingual extension and three roots, is morphologically very different from that of A. elvirae. The P2, P3 and P4 of A. elvirae are markedly smaller than those of A. adroveri (Rümke, 1985a) from Aljezar-B (typelocality), whereas the upper canine is longer; the P4 and the molars are similar in size in both species. Direct comparison of our material with that of A. adroveri from Aljezar-B stored at the university of Lyon-1 revealed several morphological differences. The P4 of A. adroveri have a better-differentiated talonid basin than that of A. elvirae. The lower molars of A. adroveri have a less-developed parastylid than those of A. elvirae. The P2 of A. adroveri from Aljezar-B is wider than that of A. elvirae and it has a small lingual lobe that is absent in the specimens from TCH-1. The protocone and the protoconule are smaller in the upper molars of A. adroveri than in those of A. elvirae. Moreover, the M2 and M3 of A. adroveri lack the accessory cusp that is present in the specimens from TCH-1. A. turolensis (Rümke, 1985a) from Masada del Valle 2 and 5, Cubla and Los Mansuetos (Teruel, Spain) has larger P2 and P2 than those of A. elvirae, whereas M1, P4 and M1 are smaller. The P4 from Masada del Valle 2 (type-locality of A. turolensis), described and figured in Rümke (1985a), have a less-developed lingual lobe than the specimens from TCH-1. We have compared our material with that of A. turolensis from Los Mansuetos kept at the university of Lyon-1. The lower molars of A. turolensis lack a parastylid; the M2 have a smaller protoconule and metaconule than the specimens of A. elvirae. The differences are more evident in the M3: A. turolensis has a very small protoconule, hardly differentiated from the protocone, and lacks an accessory cusp. There exist only three small samples ascribed to A. luteyni, from Villalba Baja 1, Masada del Valle 6 and La Fontana 1A (Teruel, Spain, Rümke, 1985a). The P4 and the molars of A. luteyni are smaller than those of A. elvirae. Morphologically, A. luteyni differs from A. elvirae by the lack of a paraconid and metaconid in its P4 and by the high connection between protocone and paracone in its P4. The premolars (except the P4) of A. elvirae from TCH-1 are notably smaller than those of A. major (Rümke, 1985a) from

Masada del Valle 7 and Valdecebro 3. The difference in size of these teeth is even more accentuated in A. verestchagini from Kosyakin Quarry (Russia), Spilia 1, Kardia and Ptolemais 1 (Greece). Nevertheless, the molars of these two species and those of A. elvirae are similar in size and M3 and M1 are even larger in the population from TCH-1. The P4 of A. elvirae is larger than that of A. major, but smaller than that of A. verestchagini. With regard to the morphology, A. major differs from A. elvirae in the absence of an accessory cusp and the smaller size of the protoconule in the upper molars and the absence of a parastylid in the lower molars. The P2 of A. verestchagini from Spilia 1 has a lingual extension and an accessory lingual root, the P4 has a less-developed parastyle and the upper molars have a smaller protoconule and metaconule than in the sample from TCH-1. The single population assigned to A. pontica (Schreuder, 1940), Polgárdi, consists of 10 measured specimens only; however, the difference in size with A. elvirae is clear: all the measured premolars of A. pontica (P1, P2, P1 and P2) are notably larger than those of A. elvirae, whereas the M2 are very similar and the M1 are smaller than those from TCH-1. The main morphological difference between these two species is the presence of a distinct lingual extension and three roots in the P2 of A. pontica. The species A. baetica Martín Suárez et al. (2001), described from Purcal-4, differs from A. elvirae in the enormous length of P2 and P4. The upper canine and the P3 are also larger in A. baetica. On the contrary, the molars and the P4 from TCH-1 and PUR-4 are very similar in size. Moreover, there are clear morphological differences between these species. The lower premolars are less molarized in A. baetica: P2 and P3 are elliptical teeth, without cristids and with very poorly developed cingulids and the P4 lack a paraconid, in contrast with those from TCH-1. The P4 of A. baetica have a proportionally smaller protocone than those of A. elvirae. The M3 of A. baetica differs from those of A. elvirae by the smaller and less prominent metaconule and the less-individualized protoconule. In A. bifida (Engesser, 1980), P3 is longer than P2; this feature distinguishes this species not only from A. elvirae, but also from the rest of the species of the genus. The P3 of A. bifida is larger than that of A. elvirae from TCH-1; on the contrary, the

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P1, P4, M1 and M2 are notably smaller. Moreover, the material from the type-locality of A. bifida (Dinar-Akçaköy) figured in Engesser (1980) and the available casts of the teeth from this locality allowed us to observe several morphological differences with A. elvirae: the I1 of A. bifida is narrower and has a much longer medial crest than that of A. elvirae; the M1 and M2 of A. bifida have a more-developed accessory cusp, the P4 has a much smaller protocone and lacks a metacone; the M1 and M2 of A. bifida lack a parastylid. One of the species that most resemble A. elvirae is A. brailloni (Rümke, 1985a). The size of some teeth (P4 and molars) is similar in the population from TCH-1 and in A. brailloni from Sète (type-population), Escorihuela, Orrios and Gorafe-2 (Rümke, 1985a). On the contrary, other dental elements, especially the lower premolars and the P2, are notably smaller in the population from TCH-1. We have compared the material of A. elvirae from TCH-1 with that of A. brailloni from TCH-1B, described later in this paper. In general terms, the P4 and the molars have similar dimensions in both populations; the P4 is similar in length, but is clearly narrower in TCH-1. The rest of the premolars are notably smaller in TCH-1 (we have not compared the size of P1, which has been not found in TCH-1B and P3, not found in TCH-1). In addition to the evident difference in size, the comparison with the populations of A. brailloni from Sète (type-locality) and Balaruc 2, kept at the university of Lyon-1, and with the material from TCH-1B, reveals several morphological differences between this species and A. elvirae. The lower premolars of A. brailloni are more molarized than those of A. elvirae: the P2 have a distinct paraconid and a narrow talonid basin that do not exist in the specimens from TCH-1. The P3 of A. brailloni are more robust and have two bulges at the anterior and posterior ends that are not present in the specimens from TCH1; on the other hand, the specimens of A. elvirae have sharper crests. The P4 from Sète, Balaruc 2 and TCH-1B are proportionally wider than those of A. elvirae and have a notably wider talonid basin. In the lower molars of A. brailloni, the parastylid is less developed than in those of A. elvirae, or even absent. The P2 from TCH-1 lack the small lingual extension that is present in the specimens of A. brailloni. The upper molars of both species are quite similar, but those from TCH-1 have a better-individualized protoconule, separated from the protocone by a deeper valley and a more developed accessory cusp than the specimens of A. brailloni from TCH1B, Sète and Balaruc 2. The teeth from TCH-1 are, in general, larger than those from Kardia, type-population of A. dekkersi (Rümke, 1985a), now included in the species A. getica (see Rzebik-Kowalska, 2005). The lengths of the P2, P3, P4, C sup. and P2 from TCH-1 fit the size ranges of those from Kardia; the mean values are similar in both localities, although somewhat higher in TCH-1 (except for the single P2 from TCH-1, slightly smaller than those from Kardia). On the other hand, the difference in size is very evident in the P4 and in the molars, which are clearly larger in A. elvirae. The P2:P3 ratio is lower in TCH-1 (1.09) than in Kardia (1.16); however, this is not an significant difference,

since only one P2 from TCH-1 was available. The P2:P3 ratio cannot be estimated in TCH-1, since no P3 has been found. Similarly, the lengths of the specimens of A. getica from Maramena and Ptolemais 3 (Rümke, 1985a) are, in general, smaller than those of A. elvirae from TCH-1; the differences in size are especially important in the P4 and the molars. The only material of A. getica from Ciupercerni is a mandible with some teeth; the P2 and P3 are similar in size to those of A. elvirae, whereas the P4 and the molars are notably smaller. There are also some morphological differences between A. elvirae from TCH-1 and A. getica. The P1 of the latter species has a unique root, whereas the specimens from TCH-1 have two fused roots. The P2 and P3 of A. getica from Kardia and Ptolemais 3 have a more rounded outline and a longer and thicker posterior cingulid than the specimens from TCH-1. The P4 of A. elvirae have a more developed paraconid and a notably narrower posterior cingulid than those of A. getica. The lower molars of A. elvirae have a very narrow posterior cingulid and a well-developed parastylid; on the contrary, those of A. getica have a broad posterior cingulid and lack a parastylid. The P4 of A. getica have neither a metacone nor a widening of the posterocrista. All the M1 and M2 from TCH-1 show a narrow metastyle, which is absent in the specimens of A. getica. The M2 of A. getica have a more triangular outline than those of A. elvirae because they have a shorter lingual face. The M3 of A. getica have a less-developed accessory cusp than those of A. elvirae. The supposed occurrence of A. getica in the Guadix-Baza Basin poses a problem. The populations from Gorafe-1 (Go-1, Rümke, 1985a; Martín Suárez, 1988) and Botardo-C (Bo-C, Martín Suárez, 1988) were originally ascribed to D. dekkersi (now A. getica, following Rzebik-Kowalska, 2005). Van den Hoek Ostende et al. (1989) considered this species as part of a lineage of the genus present in Eastern Europe; therefore, these authors indicate that the presence of ‘‘D. dekkersi’’ in Go-1 may be based on an incorrect determination due to the limited material available from this locality, or to a migration. The specific determination of the populations from Go- and Bo- is outside the scope of the present paper. Certainly, the material is so scarce that it is difficult to determine, but some biometrical features exist that point to a specific ascription different from A. getica: the M1, M3, M1 and M2 from Go-1 and Bo-C are slightly larger than those of A. getica from Kardia, Ptolemais and Maramena. Anyhow, it is evident that the population of A. elvirae from TCH-1 shows intermediate features between those from Bo-C and Go-1 and the sample of A. brailloni from TCH-1B. The size is, in general terms, larger than in Bo- and Go- and smaller than in TCH-1B, even though the size of the P1-P3 (lower and upper) from TCH-1 is more similar to that of the specimens from Go-1 and Bo-C, whereas the dimensions of the P4-M3 resemble more those of A. brailloni (Table 3). We have compared the material from TCH-1 with the few specimens from Bo-C and Go-1 stored in the university of Granada. The size is notably larger in TCH-1, especially in P4 and M1. With regard to the morphology, we have not observed any important differences, but that may be due to the scarcity of

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389

Table 3 Mean lengths (in mm) of different teeth and ratios between the mean lengths of the P2:P3 and the P2:P3 in the populations of Archaeodesmana from Western Europe Tableau 3 Longueurs moyennes (en mm) des dents et ratios de longueurs moyennes P2/P3 et P2/P3 dans les populations de Archaeodesmana de l’Europe occidentale Species

Locality

A. sp. 3 A. sp. 3 A. brailloni A. brailloni A. brailloni A. brailloni A. brailloni A. elvirae ‘‘A. brailloni’’ A. sp. 2 * A. sp. 2 ** A. sp. 1 A. baetica A. luteyni A. major A. major A. luteyni A. turolensis A. turolensis A. turolensis A. adroveri A. turolensis A. vinea A. vinea

Mo TCH-3 Ba Es TCH-1B Se Go-2 TCH-1 Or Go-1 Bo-C CLC-3 PUR-4 VB Ma-7 Va-3 Ma-6 Ma-5 LM Cu LA Ma-2 DD Ko

P2

P3

P4

M1

M2

C sup

P2

1.15

1.59

1.29 1.56 1.44 1.55 1.49

1.45 1.48 1.41

1.37 1.53 1.41

1.26 1.46 1.24

2.03

1.43

1.60 1.51

1.42 1.45 1.25 1.29 1.31 1.33 1.41 1.31 1.14 1.15

1.53 1.52 1.51 1.60 1.52 1.45 1.44

1.74 1.73 1.64 1.68

2.66 2.49 2.58 2.52

2.60 2.50 2.50 2.47

1.48 1.38 1.34

1.67 1.68 1.58 1.57 2.13 2.18 1.65 1.72 1.73

2.61 2.64

2.48 2.45 2.23

1.34 1.41 1.20

2.39 2.70 2.63 2.44 2.63 2.57

2.44 2.31 2.59 2.45

1.68 1.39 1.62 1.54

1.74

2.31

1.45

1.65 1.82 1.63 1.63 1.60

2.45 2.51 2.47 2.33 2.40

2.42 2.38 2.40 2.49 2.33 2.30 2.25

1.39 1.21 1.41

1.76 1.84 1.77 1.76 1.65 1.81 1.53 1.47 2.02 1.70 1.54 1.97 1.89 1.68 1.76 1.76 1.77 1.78 1.76 1.51 1.40

P3

P4

1.34

2.28 2.27 2.15 2.20 2.25 2.17 2.31 2.25 2.05 1.94 2.61 2.35 1.91 2.14 2.11 2.07 2.18 2.06 2.22 2.29 2.11

M1

M2

P2:P3

P2:P3

0.99 1.05 1.06

1.12 1.18 1.17 1.16

3.26

1.57 1.58 1.51 1.52 1.61 1.41 1.42 1.88 1.55 1.38 1.50 1.53 1.46 1.37 1.41 1.58 1.28 1.26 1.31

1.92

3.29 3.30 3.07 3.25 3.28 3.29 3.05 2.96 3.19 2.97 3.01 2.99 3.06 3.12 2.85 3.19 3.25 3.08 2.82

2.30 2.35 2.27 2.22 2.26 2.25 2.36

2.57 2.13

1.09 1.05 1.14

1.42

2.24

1.06 1.04

2.24

1.19 1.16 1.13 1.13 1.16 1.27 1.25

2.14 2.39 2.29 2.13 2.03

1.12 1.06 1.04 1.07 1.10 1.12 1.31 1.24 1.15 1.28 1.26 1.14 1.38 1.20 1.07

Abbreviations/Abréviations : Ko: Kohfidisch. DD: Dorn-Dürkheim. LA: Aljezar B. Ma: Masada del Valle. Cu: Cubla. LM: Los Mansuetos. Va: Valdecebro. VB: Villalba Baja 1. PUR: Purcal. CLC: Calicasas. Bo: Botardo; Go: Gorafe. Or: Orrios. Se: Sète. Es: Escorihuela. Ba: Balaruc. Mo: Moreda. The Ruscinian populations from the Guadix-Baza Basin are indicated in bold type. * Dibolia dekkersi in Rümke (1985a, 1985b) and Martín Suárez (1988); Archaeodesmana getica in van den Hoek Ostende and Furió (2005). ** Dibolia dekkersi in Martín Suárez (1988); Archaeodesmana getica in van den Hoek Ostende and Furió (2005).

the material, since many dental elements are not available in Bo-C and Go-1. The mesostyles are higher and closer to each other and the protoconule is smaller in the only M1 from Bo-C than in the specimens from TCH-1. The phylogenetic relationships of A. elvirae are not clear. Taking into account the successive increase in size in the populations from Bo-C, Go-1, TCH-1 and TCH-1B and the geographic proximity of these localities, the most logical option is to include all these populations in the same lineage. In the phylogenetic reconstructions proposed by Rümke (1985b) and van den Hoek Ostende et al. (1989), there is no phylogenetic relationship between the species dekkersi (= getica) and brailloni, which belong to the ‘‘Eastern lineage’’, on the one hand and the ‘‘Western lineage’’, on the other. So, we think that the specimens from Bo-C and Go-1, previously assigned to A. getica, correspond in fact to another species, which would be the ancestor of A. elvirae and the latter species would be the ancestor of A. brailloni. The assumption of these phylogenetic relationships involves the change of the previously proposed schemes, as we will discuss below. A. brailloni (Ru¨mke, 1985a) Fig. 2 1985a. D. brailloni n. sp. – Rümke: 153, Text-Figs. 10.2, 18.1, 24.2, 25.3, 26.2, 26.3, 27.3, 52.3, 66, 67, 68.

1986. D. brailloni Rümke – Crochet: 159, Pl. 3, Figs. 3, 8. 1986. Dibolia cf. brailloni Rümke – Crochet, 159, Pl. 3, Figs. 1, 2, 6, 7. 1988. D. brailloni Rümke – Martín Suárez, 47. 1989. D. brailloni Rümke – van den Hoek Ostende et al., 50. 1990. D. brailloni Rümke – Mein et al., 126. 1995. A. brailloni (Rümke) – Hutterer, 172. 2001. A. brailloni (Rümke) – Martín Suárez et al., 553. 2005. A. brailloni (Rümke) – Minwer-Barakat, 331, Pl. 19, Figs. 1–21. 2005. A. brailloni (Rümke) – van den Hoek Ostende and Furió, 259. Type locality: Sète (Southern France). Holotype: Skull with right I1–I2, C sup.–M3 and left I1, C sup.–P2, P4–M2, stored at the Dr. Braillon collection, Beauvais, France. Referred specimens: 2 I1 (TCH-1B 625; TCH-1B 702); 5 I2 (TCH-1B 626–630); 5 I3 (TCH-1B 631–635); 6 C inf. (TCH1B 636–641); 4 P1 (TCH-1B 642–645); 3 P2 (TCH-1B 646–648); 4 P3 (TCH-1B 649–652); 4 P4 (TCH-1B 653–656); 6 M1 (TCH1B 657–662); 6 M2 (TCH-1B 663–668); 10 M3 (TCH-1B 669– 678); 1 I1 (TCH-1B 703); 1 I2 (TCH-1B 679); 3 I3 (TCH-1B 680– 682); 3 C sup. (TCH-1B 683–685); 2 P2 (TCH-1B 686, 687); 3 P3 (TCH-1B 688–690); 1 P4 (TCH-1B 691); 1 M1 (TCH-1B 692); 5 M2 (TCH-1B 693–697); 4 M3 (TCH-1B 698–701).

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Fig. 2. A. brailloni (Rümke, 1985a) from TCH-1B. A, right I1, TCH-1B 625. B, right I2, TCH-1B 630. C, right I3, TCH-1B 634. D, right C inf., TCH-1B 638. E, right I1, TCH-1B 703. F, right I2, TCH-1B 679. G, right I3, TCH-1B 681. H, right P1, TCH-1B 644. I, right P2, TCH-1B 648. J, left P3, TCH-1B 649. K, left P4, TCH1B 653. L, left M1, TCH-1B 657. M, left M2, TCH-1B 663. N, left M3, TCH-1B 669. O, right C sup., TCH-1B 685. P, left P2, TCH-1B 686. Q, left P3, TCH-1B 688. R, left P4, TCH-1B 691. S, right M1, TCH-1B 692. T, right M2, TCH-1B 696. U, right M3, TCH-1B 700. A–E, posterior view. F–U, occlusal view. Scale bar = 1 mm.

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Occurrence in the studied area: Tollo de Chiclana-1B (TCH-1B), late Ruscinian. Measurements: See Table 1. Description: I1: The crown is relatively high; the enamel base extends equally far down anteriorly and posteriorly (morphotype ‘‘a’’ sensu Rümke, 1985a). The labial surface is flat. The posterior face is slightly convex and crossed by a wide rib from the top to the base. There is a narrow, well-marked cingulid on the posterior border. The root is long and triangular in section. I2: The crown is high, with a triangular cross-section. The apex is compressed, forming a straight cutting edge. There is a well-marked cingulid at the base of the lingual and posterior faces; this cingulid forms a small cuspid at the posterior end of the tooth. A sharp posterocristid connects the apex to the posterior cuspid. The root is long and anteroposteriorly compressed. I3: The crown is lower than in I2 and markedly asymmetrical, with a convex labial face and a slightly concave lingual face. The cingulid is more developed on the posterior face than on the lingual one. The anterocristid is short. A sharp posterocristid crosses the posterior face diagonally, from the apex of the tooth to the posterolabial corner. The root is long and anteroposteriorly compressed. C inf.: Incisor-shaped teeth, similar in size and morphology to the I3. The labial face is convex; the lingual one is slightly concave. The anterocristid is stronger than in the I3. The posterocristid is straight. The cingulid is poorly developed and restricted to the posterior border. The root is long and anteroposteriorly compressed. P1: The crown is low, with rounded outline and a pointed posterior end. The enamel base rises at the anterior part of the tooth. The labial face is convex; the lingual one is slightly concave. The cingulid is narrow and weak on the lingual part of the tooth and widens at the posterior end. The anterocristid is poorly marked. A posterocristid is absent. The root is transversely compressed; it has a deep groove on its labial side. P2: The crown is higher than in P1. The enamel base rises at the anterior and posterior ends of the tooth. The labial face is convex. The lingual face is convex in its anterior part and concave in its posterior portion. The anterolabial side is flat or slightly concave and the anterolingual end is pointed. The anterocristid and the posterocristid are sharp and well marked. The cingulid is thick and limited to the posterior border. It forms a small protuberance at the meeting point with the posterocristid. A small but distinct paraconid is present. There is a narrow talonid basin, closed posteriorly by the cingulid. There are two roots. P3: The crown is notably lower than in the P2. The outline is nearly oval, with a pointed anterior end. The enamel base rises

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at the anterior end of the tooth. The labial face is convex, although the anterolabial part may be flat or slightly concave. The lingual face is convex in its anterior part and becomes concave in its posterior portion. The anterocristid and the posterocristid are weak, or even absent in some specimens. There is a protuberance at the anterior end of the tooth, which forms a distinct paraconid in one specimen. The cingulid is thick and well marked; it extends from the posterolabial to the posterolingual part of the tooth. There is a small bulge at the intersection between the posterocristid and the cingulid. There are two divergent roots. P4: High-crowned teeth. The lingual and posterior borders are straight, the anterolabial one is curved. The posterior part is wider than the anterior one. The lingual face is nearly flat. The labial face is convex. The anterocristid is connected to a wellindividualized paraconid. A sharp posterocristid crosses the posterior face of the protoconid; it curves near the base of the crown towards the posterolingual end of the tooth, closing the talonid basin posteriorly. In one specimen, this posterolingual extension of the posterocristid is widened, forming an entostylid. The talonid basin is narrow and closed lingually by another crest, which connects the posterolingual part of the protoconid to the posterolingual end of the tooth. There is a thick, well-marked cingulid on the posterior and posterolabial borders and a narrower, poorly defined anterior cingulid. There are two roots. M1: The anterolabial cingulid is narrow; it becomes very thin or interrupted at the base of the protoconid and appears again farther backwards, occupying the base of the re-entrant valley. There is no parastylid. The oblique cristid does not reach the top of the metaconid. The posterior cingulid is narrow and connected to a medium-sized, oval entostylid. There are two roots. M2: The anterolabial cingulid is wider than in M1; it narrows at the base of the protoconid and reaches the base of the hypoconid. Contrary to M1, the cingulid broadens at its anterolingual part, forming a wide and long parastylid. The oblique cristid does not reach the top of the metaconid. The posterior cingulid is very narrow and connected to an oval, prominent entostylid. There are two roots. M3: The anterolabial cingulid is wide in its anterior part and becomes narrower at the base of the protoconid; it reaches the base of the hypoconid. This cingulid widens anterolingually, forming a parastylid. The oblique cristid is shorter than in the M1 and M2. There is no posterior cingulid. Only two out of ten specimens have a small entostylid; in some other specimens, there is a weak swelling of the enamel at the posterolingual end of the tooth. There are two roots. I1: The crown is low, with triangular cross-section. The labial face is convex, the lingual face is concave and the anterior

Fig. 2. A. brailloni (Rümke, 1985a) de TCH-1B. A, I1 droite, TCH-1B 625. B, I2 droite, TCH-1B 630. C, I3 droite, TCH-1B 634. D, C inf. droite, TCH-1B 638. E, I1 droite, TCH-1B 703. F, I2 droite, TCH-1B 679. G, I3 droite, TCH-1B 681. H, P1 droite, TCH-1B 644. I, P2 droite, TCH-1B 648. J, P3 gauche, TCH-1B 649. K, P4 gauche, TCH-1B 653. L, M1 gauche, TCH-1B 657. M, M2 gauche, TCH-1B 663. N, M3 gauche, TCH-1B 669. O, C sup. droite, TCH-1B 685. P, P2 gauche, TCH1B 686. Q, P3 gauche, TCH-1B 688. R, P4 gauche, TCH-1B 691. S, M1 droite, TCH-1B 692. T, M2 droite, TCH-1B 696. U, M3 droite, TCH-1B 700. A–E, vue postérieure. F–U, vue occlusale. Échelle = 1 mm.

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face is flat and shorter than the other two. The anterior and labial faces form a nearly right angle. There is a shallow indentation on the ventral edge of the anterior face, which divides the tooth in two lobes. The labial lobe is larger and slightly higher than the medial one. The base of the crown is damaged, so the cingulum cannot be observed. On the lingual face, the enamel forms a broad rib, which extends from the base to the middle of the crown. The medial crest is long and sharp. The root is not preserved. I2: The crown is low, conical, with elliptical outline. The anterocrista and posterocrista are sharp. There is a cingulum on the lingual border; it widens especially at the anterior, posterior and lingual ends of the tooth. The root is long and thick, with circular cross-section. I3: These teeth are smaller than the I2 and their outline is circular. The anterocrista and posterocrista are well marked. The cingulum occupies the entire lingual border and broadens at the anterior, posterior and lingual extremities. The root is long and thinner than in I2. C sup.: Conical, high-crowned teeth, with elliptical outline. The enamel base rises slightly at the middle of the labial and lingual faces. The lingual face is practically flat; the labial face is convex. There is neither an anterocrista nor a posterocrista. The anterior cingulum is well marked and restricted to the anterior border. The posterior cingulum is wider and reaches the middle of the lingual face. Two out of three specimens preserve the roots. One of them has two divergent roots; the other one has two fused roots that separate near their extremes. P2: The outline is oval. The length is notably larger than the width. The posterior part is wider than the anterior one. The anterior cingulum is thick and bulged, limited to the anterior border. The posterior cingulum is wide and extends from the posterior part of the labial face to the middle of the lingual border. At this level, the cingulum rises slightly and widens, forming a vestigial protocone, which protrudes over the outline of the tooth, forming a small lingual lobe. The enamel base rises at the middle of the labial and lingual faces. The posterocrista extends from the top of the paracone to the posterior cingulum. There are two roots; the posterior one is wider than the anterior one and it extends below the lingual lobe. P3: Low-crowned teeth, smaller and more molarized than P2. The labial border is curved; the anterior, posterior and lingual extremes are pointed. There is a small, elongated protocone, separated from the paracone by a valley. The protocone forms a lingual extension. There is a short, thick and prominent parastyle. The posterolingual cingulum is narrow, low, poorly marked and connected to the protocone. There is a short and narrow posterolabial cingulum. There is no anterocrista. The posterocrista reaches the posterior cingulum. There are two roots: the anterior one is small; the other one is wide and occupies the entire posterolingual border. P4: Strongly molarized teeth, with triangular outline. The paracone is high; its labial and anterior faces are convex, whereas the posterolingual face is flat. There is no anterocrista. The posterocrista is pronounced and connected to a poorly differentiated metacone. The protocone is large, circular and separated from the paracone by a valley. The parastyle is wide

and prominent, connected to two cingula: a narrow anterolabial cingulum that disappears at the middle of the labial face and an anterolingual cingulum that ends at the level of the paracone– protocone separation. The metastyle is much narrower and less individualized than the parastyle. The posterolabial cingulum is short and narrow. The posterolingual cingulum is thick and it reaches the base of the protocone. There are two large roots in anterior and posterior position and a smaller third one below the protocone. M1: The parastyle is rounded, prominent and connected to the protoconule by a low and wide crest. The metastyle is narrow. The protocone and the protoconule are separated by a shallow groove. Below this groove, at the base of the crown, there is a small enamel bulge. There is an accessory cusp attached to the posterior part of the protocone. The metaconule is equal in size to the protoconule. The posterior cingulum is long, narrow and attached to the metacone. There are three main roots at the lingual, anterolabial and posterolabial ends and another minuscule root below the centre of the tooth. M2: The parastyle, less prominent than in M1, is a widening of the anterolabial cingulum. The metastyle is poorly developed or absent. The protoconule is well individualized, separated from the protocone by a groove. At the base of this groove, there is a small rounded tubercle. The protocone is shifted towards anterior. The accessory cusp is large, but poorly individualized from the protocone. The metaconule is sharp and separated from the metacone and from the accessory cusp by two valleys. The anterior cingulum is narrow; the posterior one is even weaker and lower, attached to the metacone. There are three main roots in lingual, anterolabial and posterolabial position and a small secondary root in the centre of the tooth, connected to the other ones by crests. M3: The parastyle is narrow but distinct in three specimens and very much reduced in the other one. The single arm of the metacone thickens at its labial end, forming a poorly defined posterior mesostyle. The protoconule is low and very close to the protocone. There is a thick crest connected to the posterior part of the protocone, but there is no distinct accessory cusp. The metaconule is notably higher and better individualized than the protoconule and protrudes over the posterior border. The anterior cingulum is very weak; it connects the parastyle to the top of the protoconule, but at the level of the paracone it becomes very thin or even interrupted. There are two minuscule protuberances at the base of the crown on the anterior and posterior sides of the protocone. There are three main roots, in lingual, anterolabial and posterolabial position. There is a small secondary root in the centre of the tooth, connected to the main roots by three crests. Comparisons: The measurements of the specimens from TCH-1B fit the size range of the type population of A. brailloni (Sète) for almost all dental elements. The only significant differences are in the lower canine, which is considerably larger in Sète and the P1, which is larger in TCH-1B. We have to take into account, however, that these teeth are extremely scarce in the sample from Sète (only one C inf. and one P1). For the rest of the teeth, the mean lengths and widths are very similar in both populations; in general, they are slightly larger in TCH-

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1B, except for P4 and P4, which are somewhat smaller. Anyway, the differences are very subtle. The P2:P3 ratio (1.06 in Sète; 1.05 in TCH-1B) and the P2:P3 ratio (1.17 in Sète; 1.18 in TCH1B) are also very similar. We have compared the measurements with those of other populations of A. brailloni: Escorihuela, Go-2 and Orrios, all of them with few specimens (Rümke, 1985a). Lengths and widths are similar in these populations and in TCH-1B for all the teeth, except the C inf. (smaller in TCH-1B) and the P1 (notably smaller in Escorihuela). In general, the specimens from Go-2 are slightly smaller than those from TCH-1B. The differences in size with the specimens from Orrios do not show any consistent pattern; they may be due to the scarcity of the material. The dimensions of A. brailloni from Balaruc 2 (Crochet, 1986) are also very similar to those from TCH-1B, although the mean values are slightly higher in the French locality. The original definition of A. brailloni does not include any description of the molars (it contains the description of the rest of the teeth) and the figured material from the type population (Sète) is scarce. Nevertheless, the direct comparison of the material from TCH-1B with the teeth and casts of A. brailloni from Sète and Balaruc 2 kept in the university of Lyon-1 has allowed us to prove that there are no significant morphological differences between these populations. The molars (upper and lower) from TCH-1B are morphologically identical to those from the French localities. The differences in the premolars are very subtle. For example, the P4 from Sète and Balaruc have a lingually open talonid basin, whereas in the specimens from TCH-1B the posterolingual cristid closes the talonid basin. The population from TCH-1B presents clear biometrical differences with most species of Archaeodesmana. All the dental elements from TCH-1B are clearly larger than those of A. getica from Kardia, Maramena and Ptolemais 3 (Rümke, 1985a). The P4 of A. brailloni from TCH-1B is shorter than that of A. adroveri and the upper canine is longer. The population from TCH-1B can be distinguished from A. baetica because in the latter the length of P2, P4 and the upper canine is much larger. The teeth from TCH-1B are larger than those of A. luteyni, especially the upper premolars. A. vinea has a much higher P2:P3 ratio, and a notably smaller size than A. brailloni from TCH-1B. The P2 and the upper canine are clearly larger in A. major than in the population from TCH-1B and the P2:P3 ratio is higher. A. brailloni from TCH-1B is much smaller than A. verestchagini. A. brailloni from TCH-1B is markedly larger than A. bifida; the P4 from TCH-1B have a more developed protocone and metacone than those of A. bifida. The population from TCH-1B differs from A. pontica in morphological features such as the number of roots of the P1, P2 and P3. A. turolensis differs from the population from TCH-1B by its smaller size and by the two roots of P1; moreover, the P3 of A. turolensis lacks a protocone and in P4 this cusp is less developed than in A. brailloni. The upper molars from TCH-1B have better-individualized lingual cusps than those of A. turolensis. A. brailloni from TCH-1B is clearly larger than Archaeodesmana sp. from Go-1 and Bo-C, previously ascribed to

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Fig. 3. Archaeodesmana sp. from TCH-3. A, right P4, TCH-3 233. B, right P3, TCH-3 232. A, B, occlusal view. Scale bar = 1 mm. Fig. 3. Archaeodesmana sp. de TCH-3. A, P4 droite, TCH-3 233. B, P3 droite, TCH-3 232. A, B, vue occlusale. Échelle = 1 mm.

A. getica and here, considered indeterminate at species level. With regard to A. elvirae from TCH-1, the differences are mentioned in the comparisons of that species; summarizing, the lower premolars, the upper canine and the P2 are notably larger in A. brailloni from TCH-1B and the lower premolars are more molarized. A. brailloni is the most recent species of the genus documented in Western Europe, although populations without a specific ascription exist in younger localities, like Moreda (Rümke, 1985a) and TCH-3, described below. Archaeodesmana sp. Fig. 3 Referred specimens: 1 P3 (TCH-3 229); 1 M2 (TCH-3 230); 1 M3 (TCH-3 231); 1 I1 (TCH-3 351); 1 P3 (TCH-3 232); 1 P4 (TCH-3 233); 1 M2–3 (TCH-3 234). Occurrence in the studied area: Tollo de Chiclana-3 (TCH-3), early Villafranchian. Measurements: Table 1. Description: P3: The crown is relatively high; the outline is elliptical, with the anterior end pointed. The enamel base rises at the anterior part of the tooth. The labial face is convex. The lingual face is convex in its anterior part and slightly concave posteriorly. The anterocristid and the posterocristid are poorly marked. The cingulid is thick and well defined; it occupies the entire lingual border and it has a small bulge at the meeting point with the posterocristid. There is one single root. M2: The only specimen is incomplete; it corresponds to the talonid and the posterior part of the trigonid. The oblique cristid reaches the top of the metaconid. The posterior cingulid is short, straight and narrow. There is an entostylid. M3: The anterolabial cingulid is narrow, but it broadens markedly at the anterolingual end, forming a very wide and prominent parastylid. The oblique cristid reaches the top of the metaconid. There is neither a posterior cingulid nor an entostylid. The roots are not preserved. I1: The tooth is damaged in its posterior part. The labial face forms a straight angle with the anterior one. The indentation of the edge of the anterior face is deep and displaced towards the medial part, so the tooth is divided into two lobes of different size. The medial lobe is narrow and it has a sharp apex. The

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labial lobe is similar in height to the medial one, but much broader; it forms a cutting edge backwards. The medial face is short; it has a triangular enamel-free area at its base (morphotype ‘‘AN’’ of Rümke, 1985a). The medial crest is long and sharp. The root is not preserved. P3: The crown is relatively high. The outline is rounded, with an extension on the lingual part. There is a low, poorly differentiated protocone, connected to the paracone by a small crest. The parastyle is large, rounded and very prominent. The posterocrista is well marked; it reaches the base of the crown and is connected to the metastyle. The metastyle is smaller and less prominent than the parastyle. The posterolingual cingulum is low and narrow; it connects the metastyle to the base of the protocone. There are two roots. P4: Large, strongly molarized teeth, with triangular outline. The labial face of the paracone is convex and its lingual face is concave. The parastyle is partly broken; it is unusually large, round and very prominent. It is connected to the paracone by the anterocrista, which attains one third of the height of the crown. The posterocrista is marked; it connects the paracone to a poorly individualized metacone. The lingual extension is very wide. The protocone is separated from the paracone by a deep valley. The posterolingual cingulum is marked and high; it reaches the base of the protocone, closing a posterolingual valley. There is a short and thick anterolingual cingulum between the parastyle and the lingual extension. There are two short, but well-differentiated cingula, anterolabial and anterolingual, which delimit two small basins at the base of the paracone. There are three roots: the anterior root is large; the posterior and lingual ones are fused. M2–3: The only fragment corresponds to the lingual part of a second or a third upper molar. The protoconule is separated from the protocone by a very deep valley. There is a long and thick posterior crest connected to the protocone, but there is no a distinct accessory cusp. The metaconule is smaller and lower than the protoconule. Comparisons: The occurrence of a short I1 with a bifid tip confirms that the Desmaninae from TCH-3 belongs to the genus Archaeodesmana. The scarcity of material makes a specific determination impossible. However, we have compared the measurements with those of the different species of the genus that existed during the Pliocene. The P3 and the P3 of Archaeodesmana sp. from TCH-3 are clearly smaller than those of A. brailloni, whereas the P4 is similar in size. On the contrary, the measurements of P3, M3 and P3 from TCH-3 fit the size range of the population of A. getica from Kardia and the P4 is clearly smaller than that of the latter species. The P3 of A. bifida is much longer than the specimen from TCH-3. All the specimens are smaller than their equivalents in A. baetica from Purcal-4. Rümke (1985a) mentioned the existence of an assemblage of only nine teeth of Archaeodesmana in Moreda (Southern Spain), characterized by their small size. Unfortunately, the few measured specimens from Moreda (P1, C sup., P1, P2, M1 and M2) do not coincide with the elements found in TCH-3, so we cannot compare the dimensions of any dental element. Nevertheless, the specimens from both localities are relatively

small. On the other hand, the morphology of the I1 from TCH-3 resembles that of the specimen from Moreda, with a deep indentation on the anterior border, a high and narrow medial lobe and a high enamel free area on the medial side. This similarity, added to the geographic proximity of the two localities and their equivalent age, indicate that these populations may belong to the same species. Although she did not make a specific determination, Rümke (1985a) indicated that the material from Moreda could correspond to D. dekkersi (= A. getica), because of the resemblance in size with that species. We think this suggestion is incorrect. The citations of ‘‘D. dekkersi’’ (A. getica in van den Hoek Ostende and Furió, 2005) in the Iberian Peninsula are here considered to be incorrect. Anyway, these citations correspond to upper Turolian (Bo-C, Martín Suárez, 1988; Nívar, Curva, Cacín-1, Ruiz Bustos, 2002) and lower Ruscinian (Go-1, Rümke, 1985a; Go-4, Martín Suárez, 1988) localities. The only species of the genus found in the upper Ruscinian in Western Europe are A. elvirae from TCH-1 and A. brailloni, identified in the Guadix-Baza Basin at Go-2 (Rümke, 1985a) and TCH-1B and also in several localities from the Teruel region (Rümke, 1985a; Mein et al., 1990; van den Hoek Ostende and Furió, 2005). These data suggest a phylogenetic relationship between the species A. brailloni and the populations from the lower Villafranchian sites of Moreda and TCH-3; however, more abundant material would be necessary to clarify the relationships between Archaeodesmana sp. from Moreda and TCH-3 and the rest of the species of the genus. 5. New data on the phylogeny of the ‘‘Western European lineage’’ of Archaeodesmana The study of fossil Desmaninae and more specifically, of Archaeodesmana is problematic mainly because of the scarcity of published data. The classification of the genera and species of Desmaninae proposed by Rümke (1985a) is practically the only reference for the determination of the fossil representatives of this subfamily. The distinction of the various genera is clear; it is based on morphological and biometrical criteria, mainly on the morphology of the incisors, as well as on significant differences in the size of the whole dentition. On the other hand, the distinction of the species of Archaeodesmana is established by means of biometrical criteria, which are rather rigid. In fact, the differential diagnoses of the six species defined by Rümke (1985a) are based exclusively on biometrical criteria. Previous interpretations of the phylogenetic relationships between the different species of Desmaninae (Rümke, 1985b; van den Hoek Ostende et al., 1989) are also based exclusively on biometrical criteria. Besides, these interpretations were made on the basis of a rather discontinuous record that possibly does not allow to observe the changes in size and morphology clearly. For example, Rümke (1985b) noted that the ratio of the lengths of the P2:P3 diminishes from older to younger populations of Archaeodesmana; nevertheless, the finding of new populations like that of A. baetica from the lower Ruscinian locality of Purcal-4, with a much higher P2:P3 ratio

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than any other species, proves that the decrease of this parameter through time is not consistent. The discovery of new populations (those described in this paper and other populations recently found in Southern Spain) may modify the previous phylogenetic schemes. More specifically, the ‘‘Western European lineage’’ proposed by Rümke (1985a), which includes A. vinea, A. turolensis, A. major and A. brailloni, must be reconsidered in the light of the later findings. First, the occurrence of A. baetica in the locality of Purcal-4 (lowermost Ruscinian) in the Granada Basin (Southern Spain), with biometrical and morphological features that are very different from the rest of the species, evidences that the phylogeny of the genus is not as simple as previously supposed. In fact, the origin of A. baetica cannot be determined, since some features (large size of the P2 and P4; upper canine similar in size to the P2; poorly molarized premolars) are very different from those of the rest of species included in the Western lineage as proposed by Rümke. Another problem arises when we analyze the population of Archaeodesmana from the locality of Calicasas-3 (Archaeodesmana sp. 1 in Fig. 4), also situated in the Granada Basin (material under study, stored in the University of Granada). The specimens from this lower Ruscinian level are larger than those of the rest of the species of the genus (see Table 3). The measurements are only similar (though slightly larger) to those of A. verestchagini, a species only recorded from Eastern Europe. The difference in size with all the populations described from the Iberian Peninsula is highly significant. Apart from the obvious difference in size, the population from Calicasas shows several morphological features that have not been observed in any other Iberian population (the lack of talonid of P4, the asymmetry between the anterior and posterior parts of P3 and the great development of the protocone in P4). Calicasas-3 is slightly younger than Purcal-4 and the two localities are geographically close. In addition, these populations share some morphological characters like the poorly molarized P4, with a minuscule paraconid and the very reduced or absent accessory cusp in the upper molars. Therefore, Archaeodesmana sp. 1 from Calicasas-3 appears to be related with A. baetica. On the contrary, the materials from Purcal-4 and Calicasas-3 differ enormously from other populations of similar age from Southern Spain, (for example, Bo-C and Go-1 in the Guadix-Baza Basin). So, there were at least two different lineages of Archaeodesmana in the South of the Iberian Peninsula during the early Ruscinian. The populations from Bo-C and Go-1 were assigned to D. dekkersi in previous papers (Rümke, 1985a; Martín Suárez, 1988) and, therefore, excluded from the ‘‘Western lineage’’ in spite of their geographic situation. In the present paper, we consider that these populations correspond to another species (as hinted by van den Hoek Ostende et al., 1989), called Archaeodesmana sp. 2 in Fig. 4. Their phylogenetic relationships constitute another unresolved problem in the phylogeny of the genus. In this context, the population of A. elvirae from TCH-1, with intermediate characteristics between the populations from

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Bo-C and Go-1 and A. brailloni from TCH-1B, suggests the inclusion of all these populations in one single lineage: as we argued in previous paragraphs, there is a continuous increase in size of all the dental elements from the oldest to the youngest population (Fig. 5). This fact and the geographic and temporal proximity of the mentioned localities, whose biostratigraphical position is well established, indicate that the species present in Go-1 and Bo-C is the ancestor of A. elvirae from TCH-1 and this latter species is the ancestor of A. brailloni (Fig. 4). Accepting this relationship implies certain incompatibilities with previous evolutionary schemes. First, the ancestor– descendent relationship between A. major and A. brailloni would become invalid, if we accept A. elvirae as the ancestor of A. brailloni. In fact, on the basis of size, the origin of A. brailloni could be in both A. major and A. elvirae: the dimensions of some teeth (P2, P3, P4) are more similar to those of A. major, whereas the size of other elements (C sup., P2, P4, M1) is more similar to that of A. elvirae. The morphology of the upper molars of A. brailloni resembles more that of A. elvirae than that of A. major: the latter species has a notably lessdeveloped protoconule than A. brailloni and A. elvirae and it lacks the accessory cusp that is present in the other species. This suggests the origin of A. brailloni to be in A. elvirae. The origin of A. elvirae is in Archaeodesmana sp. 2 from Bo-C and Go-1. The relationship between the Miocene species A. major and the continuous lineage proposed in this paper, formed by A. sp. 2 A. elvirae - A. brailloni, is not clear. Besides, the presence of A. brailloni in the lower Ruscinian site of Orrios (Rümke, 1985a) must be reconsidered. The size of the specimens from Orrios resembles that of A. brailloni, but also that of A. major, a species found in older localities from the Teruel region (Masada del Valle 7, Valdecebro 3, upper Turolian). With respect to the morphology, there are some differences between the material from Orrios and other populations assigned to A. brailloni: the premolars from Orrios are, in general, more molarized; the lower premolars have a better developed paraconid and the upper premolars have a more pronounced lingual extension and a better developed protocone. Moreover, the locality of Orrios (early Ruscinian, MN14) is notably older than the rest of localities from which A. brailloni has been reported (Sète, Go-2, TCH-1B, late Ruscinian; Escorihuela and Balaruc, early Villafranchian). None of the species of Archaeodesmana has such a long temporal distribution. In addition, no data are available on any other population of A. brailloni from the early Ruscinian; there are citations of its presence in several localities in the area of Teruel (Mein et al., 1990), but neither the descriptions nor the measurements of the teeth have been published. The specific determination of the material from Orrios exceeds the scope of this paper but, for all the exposed reasons, we consider the previous assignment not to be conclusive (this population is quoted as ‘‘A. brailloni’’ in Table 3). Finally, the relationships between the few specimens found in the early Villafranchian sites of Moreda (Rümke, 1985a) and TCH-3 (Archaeodesmana sp. 3 in Fig. 4) and the other species of the genus cannot be established clearly, mainly due to the scarcity of material. Rümke (1985a) mentioned the possible

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Fig. 4. Chronological distribution and phylogenetic relationships of the populations of Archaeodesmana from Southern Spain (Granada and Guadix-Baza Basins). Question marks indicate doubtful relationships. Fig. 4. Distribution chronologique et relations phylogénétiques des populations de Archaeodesmana du Sud de l’Espagne (Bassins de Grenade et Guadix-Baza). Relations douteuses indiquées par points d’interrogation.

relation between the population from Moreda and D. dekkersi (= A. getica). As we have explained, this species is present exclusively in Eastern Europe, so this relation seems to be incorrect. The most likely possibility is the relation with A. brailloni, since this is the youngest species of the genus and it has been cited in localities in the same area (TCH-1B and Go-2). In addition, although the dimensions of the P2, the upper canine, P2 and P3 from TCH-3 and Moreda are clearly smaller than those of the specimens of A. brailloni from TCH-1B (the nearest locality in space and time to Moreda and TCH-3), the size of the P4 and the M1 is similar. However, confirmation of a relationship

between A. brailloni and the populations from Moreda and TCH3 is impossible due to the scarcity of material of these populations. Table 3 shows the mean lengths of several teeth and the ratios between the lengths of P2:P3 and P2:P3 in the populations of Archaeodesmana from Western Europe, biostratigraphically ordered. In addition to the populations studied in this paper, we have represented data of the species of the ‘‘Western lineage’’ of Rümke (A. vinea, A. turolensis, A. major and A. brailloni), of other species not included in this lineage, but present in Western Europe (A. adroveri, A. luteyni, A. baetica), as well as the

Fig. 5. Ranges of the length of the premolars and molars of Archaeodesmana sp. 2 from Bo-C and Go-1, A. elvirae from TCH-1 and A. brailloni from Go-2 and TCH1B. Data of Go from Rümke (1985a); data of Bo from Martín Suárez (1988); data of TCH-1 and 1B from this paper. Fig. 5. Intervalles des longueurs des prémolaires et molaires de Archaeodesmana sp. 2 de Bo-C et Go-1, A. elvirae de TCH-1 et A. brailloni de Go-2 et TCH-1B. Données de Go d’après Rümke (1985a) ; Bo d’après Martín Suárez (1988) ; TCH-1 et 1B dans ce travail.

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measurements of populations without a specific ascription (‘‘A. sp. 1’’ from Calicasas-3, ‘‘A. sp. 3’’ from Moreda), or with a specific determination considered incorrect in the present paper (‘‘D. dekkersi’’ from Go-1 and Bo-C, named ‘‘A. sp. 2’’ in Table 3). Relatively continuous changes in the size of the whole dentition can be observed only in populations from restricted geographic areas and for certain time intervals. Such is the case of the Ruscinian populations from the Guadix-Baza Basin (BoC, Go-1, Tollo de Chiclana-1, Go-2 and Tollo de Chiclana-1B, represented in bold type in Table 3). In these populations, a clear increase in the size of the teeth can be noted from the oldest to the youngest population (Fig. 5). 6. Conclusions The phylogenetic relationships between the different species of Archaeodesmana are still unclear. In this paper, we do not claim to clarify all these relationships, but only those of the new populations here described, pointing out, however, that the phylogeny of the genus Archaeodesmana is much more complex than previously considered. Therefore, an exhaustive revision of the distinct populations of Archaeodesmana and their phylogenetic relationships would be necessary, taking into account not only the size of the teeth, but also the dental morphology, the geographic situation of the localities and the age of the fossil assemblages. We propose the phylogenetic relationship between the unnamed species from Go-1 and Bo-C, A. elvirae and A. brailloni as an alternative to the lineage previously proposed by other authors (Rümke, 1985a, 1985b; van den Hoek Ostende et al., 1989). Probably, the forms reported in Moreda (Rümke, 1985a) and TCH-3 represent the youngest species of this lineage. At least, one other lineage of this genus is present in Southern Spain during the Pliocene, formed by A. baetica and the unnamed species from Calicasas-3. Acknowledgements We thank Dr. P. Mein, who put the material stored in the University of Lyon-1 at our disposal. Careful reviews by R. Ziegler and L. van den Hoek Ostende greatly improved the manuscript. This study was supported by 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 and the FEI Quanta 400 Scanning Electron Microscope of the ‘‘Centro Andaluz de Medio Ambiente’’, Granada. References Crochet, J.Y., 1986. Insectivores Pliocènes du Sud de la France (LanguedocRoussillon) et du Nord-Est de l’Espagne. Palaeovertebrata 16, 145–171. Engesser, B., 1980. Insectivora und Chiroptera (Mammalia) aus dem Neogen der Türkei. Schweizerische Paläontologische Abhandlungen 102, 45–149. Fernández, J., Viseras, C., Soria, J.M., 1996. Pliocene–Pleistocene continental infilling of the Granada and Guadix basins (Betic Cordillera, Spain): the influence of allocyclic and autocyclic processes on the resultant stratigra-

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