concerned deposits are particularly rich in modern type, close to still extant selachian forms. This is confirmed by the presence of different species of Dasyatis. From the thermic point of view, Alvalade basin marine fishes point out to a decrease in the prevailing thermic conditions. tale. Elle est rep valade, Portugal méridional). De n rmes). Plusieurs genre qui suggère d des prédominaient, moins qu’au Torton tatifs des taxons en Geobios 39 (2006) 747–755 cause entre les gisements qui ont été exploités sont différents, ce qui est en rapport avec des conditions paléoenvironnementales. Du point de vue bathymétrique, les D * Corresponding aut E-mail addresses: 0016-6995/$ - see fron doi:10.1016/j.geobios. pératures prédominantes des eaux. Les Dasyatis sont encore très nombreux, ce qui est en contraste avec es eaux tempérées à froides, ou plus profondes et par conséquent plus froides. Des eaux modérément chau ien de Lisbonne, mais plus que dans la situation actuelle près des côtes portugaises. Les rapports quanti formes ont été reconnues. Les dépôts en question sont particulièrement riches en sélaciens de type moderne avec d’étroites affinités vis-à-vis d’espèces actuelles, ce que confirme la présence de plusieurs espèces de Dasyatis. La faune de poissons marins du basin d’Alvalade indique une diminution des tem la rareté de Raja, cerne la seule faune ichthyologique du Miocène terminal reconnue jusqu’à présent au Portugal et peut-être en résentée par plus de 10 000 dents et autres restes récoltés dans la Formation de Esbarrondadoiro (bassin d’Al ombreuses dents, décrites ici, ont été attribuées à la famille des Dasyatidae (46 % environ parmi les Myliobatifo The still abundant Dasyatis are in contrast to the rarity of Raja, a genus which suggests temperate or cold, or deeper hence colder waters. Moderately warm water conditions predominated, less than in Tortonian of Lisbon, more than exist today near the Portuguese coasts. The occurrence ratios of the taxa concerned among the exploited sites are quantitatively different. This points out to different paleoenvironmental conditions. The Dasyatis had a litoral/neritic bathymetric distribution and a benthic way of life. However, the Esbarrondadoiro pattern is very different from those from Santa Margarida and Vale de Zebro, which are nearly identical between them as far as Dasyatis gr. gigas and Dasyatis sp. are concerned: Dasyatis gr. gigas may suggest less warm waters, and this is very much compatible with other data that indicate a deeper environment and hence colder waters for Esbarrondadoiro. © 2006 Elsevier Masson SAS. All rights reserved. Résumé Ce travail con Europe occiden- Original article Latest Miocene Dasyatidae (Neoselachii, Batomorphii) from the Alvalade Basin, Portugal Dasyatidae (Neoselachii, Batomorphii) du Miocène terminal du Bassin d’Alvalade, Portugal Ausenda Cáceres Balbinoa,b, Miguel Telles Antunesa,c,* aCentro de Estudos Geológicos da Universidade Nova de Lisboa, Quinta da Torre, 2825-114 Caparica, Portugal bDepartamento de Geociências, Universidade de Évora, Apartado 94, 7002-554 Évora, Portugal cAcademia das Ciências de Lisboa, Rua da Academia das Ciências, 19, 1249-122 Lisboa, Portugal Received 13 April 2005; accepted 22 July 2005 Available online 17 August 2006 Abstract This paper deals with the only latest Miocene fish fauna so far recognized in Portugal and probably in western Europe through more than 10,000 teeth and many other remnants collected in the Latest Miocene, Esbarrondadoiro Formation (Alvalade basin, southern Portugal). A large number of teeth reported to the family Dasyatidae (ca. 46% among the Myliobatiforms) are described. Several forms were recognized. The http://france.elsevier.com/direct/GEOBIO/ asyatis fréquentent des milieux littoraux / néritiques et sont benthiques. Toutefois, le cas d’Esbarrondadoiro est très différent hor.
[email protected],
[email protected] (M.T. Antunes). t matter © 2006 Elsevier Masson SAS. All rights reserved. 2005.07.002 entiques entre eux) en ce qui concerne Dasyatis gr. gigas et Dasyatis sp. : ut à fait compatible avec d’autres données qui indiquent des milieux plus o. In the crushing type, the crown is generally spherical with a transverse crest separating the lingual and labial faces. The lin- gual face is smooth and superposed on the root through an oblique visor. The labial face is more developed, convex, and flat or with a median depression; it may be smooth, tubercular or pitted. In the clutching type, the crest is lingually developed and forms a larger or smaller cusp according to species and sex. In the grinding type, the crown is high and presents a hex- agonal to lozenge-shaped outline with well-marked marginal angles. The transverse crest is weak or absent. In some living Dasyatidae such as Himantura uarnak For- sskal, 1775, with crushing type dentition, there are some differ- ences in the arrangement of the dental files between the max- illaries and the mandible: the maxillaries present a symphysial, / Geobios 39 (2006) 747–755 lophodon Cappetta, 1980 are reported to the same family (Cap- petta, 1987). The Dasyatidae are coastal, shallow, benthic tropical and subtropical water dwellers. Some may be found in estuaries and even live in fresh water. In the base of the tail, the Dasyatidae possess a pointed and saw-toothed spine connected to a poison gland. They prey on molluscs and crustaceans. The dentition is of crushing, clutching or grinding types, whose definitions have been given by Cappetta (1970, 1987). visor; 7, labial face of the crown; 8, labial zone of the crown; 9, median labial hollow; 10, marginal angles (lateral angles); 11, lingual zone of the crown; 12, lingual marginal face of the crown; 13, lingual visor; 14, root lobe; 15, lower part of labial visor; 16, labial face of the root; 17, central foramen; 18, basal face of the root; 19, lower part of lingual visor. Fig. 1. Terminologie d’une dent de Dasyatis sp. : A, face orale ; B, face basilaire (d’après Cappetta, 1987). 1, l’échancrure linguale de la racine ; 2, lobe mésial de la racine ; 3, arête médio-linguale ; 4, face linguale de la couronne ; 5, carène transverse ; 6, visière labiale ; 7, face labiale de la couronne ; 8, zone labiale de la couronne ; 9, dépression médio-labiale ; 10, angles marginaux (angles latéraux) ; 11, zone linguale de la couronne ; 12, face marginal linguale de celui de Santa Margarida et Vale de Zebro (lesquels sont presque id Dasyatis gr. gigas peut suggérer des eaux moins chaudes, ce qui est to profonds et par conséquent des eaux plus froides pour Esbarrondadoir © 2006 Elsevier Masson SAS. All rights reserved. Keywords: Batomorphii; Dasyatidae; Alvalade Basin; Latest Miocene Mots clés: Batomorphii; Dasyatidae; Bassin d’Alvalade; Miocène terminal 1. Introduction The Alvalade basin, Esbarrondadoiro Formation (Latest Miocene) in southern Portugal yielded, at Esbarrondadoiro, Santa Margarida and Vale de Zebro, remnants from a rich and varied selachian fauna, which is represented by about 10,000 teeth (Balbino, 1995). The orders Hexanchiformes, Squaliformes, Pristiophoriformes, Squatiniformes, Lamni- formes, Carcharhiniformes, Rajiformes, Torpediniformes and Myliobatiformes are represented (Antunes et al., 1999). Several species from the genus Dasyatis are represented by 1.578 teeth as a whole, collected at the above sites. This is the only latest Miocene fish fauna so far recognized in Portugal. Its Turolian age of about ca. 5.5 Ma has been ascertained after associated, probably drifted small mammals (Antunes and Mein, 1995). The concerned stratigraphic units may thus be approximately correlated to the Messinian stage. 2. Systematics Superfamily DASYATOIDEA Whitley, 1940 Family DASYATIDAE Jordan, 1888 (Late Cretaceous - Recent) This family comprises the following genera, all with living representatives: Dasyatis, Himantura, Taeniura, Urogymnus and Urolophoides (Bigelow and Schroeder, 1953). See Com- pagno and Roberts (1984: 286). Hypolophus has often been regarded as a Dasyatis subge- nus. Cappetta (1980) revalidated Hypolophus as a genus for the extant type species sephen that was ascribed to Dasyatis. On the other hand, Nishida (1990) regarded Hypolophus as a syno- nym of Dasyatis. He also excluded Urogymnus and Urolo- phoides from the Dasyatidae, but includes Potamotrygon in this family. The extinct genera Hypolophites Strömer, 1910 and Hypo- A.C. Balbino, M.T. Antunes748 For teeth descriptions, nomenclature is presented in Fig. 1. Teeth are small, about a few millimeters only. convex area with 3–5 files of large teeth surrounded by a depressed area with small teeth, followed by another convex one similar to the symphysial area; even more distally, the teeth became smaller and smaller and became more dissymme- trical towards the commissure. Fig. 1. Tooth terminology of Dasyatis sp.: A, occlusal view; B, basal view (after Cappetta, 1987). 1, lingual notch of the root; 2, mesial root lobe; 3, median lingual ridge; 4, lingual face of the crown; 5, transverse crest; 6, labial de la couronne ; 13, visière linguale ; 14, lobe de la racine ; 15, rebord inférieur de la visière labiale ; 16, face labiale de la racine ; 17, foramen central ; 18, face basilaire de la racine ; 19, rebord inférieur de la visière linguale. / G In the mandible, the depressed symphysial area is sur- rounded by a convex area. The size and morphology of the teeth are constant except at the commissure, where teeth are small and dissymmetrical. The dental schema changes from one species to another. Comparison material from the authors own collections has been used along with collections kept at the Laboratoire de Paléontologie/ISEM–Université de Montpellier II. Genus Dasyatis Rafinesque, 1810. (Late Cretaceous - Recent) Type species: Dasyatis ujo Rafinesque, 1810. According to Bigelow and Schroeder (1953), the genus Dasyatis comprises about 30 living species (Portuguese trivial name: uje). The family Dasyatidae comprises many living genera with a similar dental morphology. Dental differences between species from the same genus are often more important than between species from different genera. This may explain why many fos- sil species that had been ascribed to Dasyatis were later reclas- sified otherwise. In Dasyatis, teeth are small, a few millimeters only. Their morphology is very typical: the crown shows, more or less in its middle part, a transverse crest that separates the labial and lingual surfaces. The labial surface has a more or less developed sub triangu- lar, middle-labial depression; right opposite there is an area that is ornamented in its upper part and smooth in its lower part, i.e. the labial area. The crown’s labial visor is the part that forms the extremity of the labial surface. The lingual surface is often divided into two marginal- lingual areas by a more or less marked median lingual ridge. The crown’s lingual visor is always narrower than the labial one and limits the lower portion of the lingual surface. Each side of the tooth shows a trihedral angle (marginal angle) where the transverse crest joins the labial and lingual visors. The genus Dasyatis has been ascribed to the Miocene of Portugal by Jonet (1968, 1978). Serralheiro (1954) ascribed to Raja and Trygon some forms that were reported to Dasyatis by Cappetta (1970). Dasyatis gr. centroura Mitchill, 1815. Figs. 2(3–5) and 3(1, 2) Synonymy: None as far as Portugal is concerned. Material: 532 teeth. Localities: Santa Margarida (96), Esbarrondadoiro (385) and Vale de Zebro (51). Description: The dissymmetrical teeth are longer than wide. The more or less spherical crown presents a wide, transverse and slightly bulged crest. Its labial surface shows a sub circular depression that is limited on the labial side by a wide external area. This depression presents an irregularly reticulate orna- mentation. The labial area is also pitted. The lower edge of A.C. Balbino, M.T. Antunes the visor is wide and shows a median protuberance. The mar- ginal angles are well marked. The median lingual ridge is a cutting one in its middle part. The marginal-lingual surfaces are depressed. Although the lin- gual visor outline is convex, in some specimens it shows a concavity in its middle part. The crown’s lingual visor is clearly over the median sulcus of the root and so it partly con- ceals the root’s lobes. The root is lingually arched, its sulcus being wide and deep. The lobes present a flat or slightly bulged basal surface and are longer in a labial-lingual sense. There is a foramen in the mid- dle of the sulcus. Lateral teeth are mesio-distally longer. Mar- ginal angles are acute. The transverse crest is sharp near the lateral angles. In male teeth, the transverse crest of the crown develops into a long, pointed cusp. The cusp presents a slight longitudi- nal depression that may be pitted. The labial surface is concave in its middle portion when seen in profile. The median-lingual crest is narrow and concave. The labial visor of the crown is very much developed. On the other hand, the root is similar to that of the female teeth. Some much smaller but closely similar teeth probably belong to young individuals of the same species. Discussion: By the presence of the crown’s middle-lingual depression as well as by their ornamentation the fossil teeth are similar to those from D. cavernosa Probst, 1877 and D. probsti Cappetta, 1970 (Cappetta, 1970: pp. 88–91, Pl. 22, Figs. 1–13 and Pl. 21, Figs. 15–23). However, these teeth differ in other characteristics: larger size, wider transverse crest, more marked marginal angles, more developed lower edge of the labial visor, convex crown lingual visor with a middle depression, and more developed root lobes. The closest similarities to the fossil teeth are those of the living species D. centroura. As far as male teeth are concerned, the fossil form has teeth with cusps that also exist in D. centroura. We think that the fossil species can be included in the D. centroura group because the general teeth morphol- ogy is similar. Dasyatis gr. gigas Günther, 1870 Fig. 2(1, 2) Synonymy: None as far as Portugal is concerned. Material: 440 teeth. Localities: Santa Margarida (118), Esbarrondadoiro (260) and Vale de Zebro (62). Description: Teeth are rather large. In occlusal view, the sometimes ornamented crown’s labial surface has a lozenge- shaped profile. The marginal angles are very distinct. The well-marked transverse crest is a cutting one, even if it is broad and truncated in some specimens. The wavy labial visor presents a broad, bulged lower edge, and a distinct mid- dle swelling. The lingual surface is concave; there is no median lingual ridge. The lingual visor profile is strongly convex and attains the sulcus of the root, which has two elliptical-profiled, labial-lingually elongated lobes. The basal surface is slightly eobios 39 (2006) 747–755 749 convex. The broad and deep middle sulcus of the root presents a central foramen, plus one or two para-central foramina. A.C. Balbino, M.T. Antunes / Geobios 39 (2006) 747–755750 The more lateral teeth are transversely elongated. The edge of the lingual visor of the crown nearly conceals the lobes of the root. Teeth reported to males are high-crowned, the crown being The more lateral teeth are broader than high. The crown is elliptical and displays an identical, pitted ornamentation. The low, concave-profiled lingual surface has no median lingual ridge. The edge of the crown’s lingual visor overlaps the root b, p ior t ; b A.C. Balbino, M.T. Antunes / Geobios 39 (2006) 747–755 751 mesio-distally compressed. The strongly pointed transverse crest is indeed a cusp with a longitudinal depression that may be ornamented. Marginal angles are distinct. The lower edge of the labial visor is broad and shows a swelling in its middle part. The wavy edge of the lingual visor is strongly convex near the root sulcus. The lobes of the root are less developed than in female teeth but their morphology is similar. Discussion: The fossil teeth are similar to those from D. rugosa Probst, 1877. However, differences in the size of the teeth, as well as in the shape of the crown, of the transverse crest and of the lingual visor, exclude attribution to D. rugosa. The teeth under study more closely resemble those from the living D. gigas. Nevertheless the crown’s ornamentation, which is a constant character in D. gigas, is lacking in most of the fossil teeth. However, this may possibly result from abrasion. Furthermore, in male teeth the transverse crest also differs, as it is pointed in the fossil ones and rounded in D. gigas. Summing up: taking into account the prevailing similarities, we think the fossil teeth can be ascribed to a species from the D. gigas group. Dasyatis pastinaca (Linnaeus, 1758) or Dasyatis marmorata Steindachner, 1892 Figs. 3(3–6) and 4(1) Synonymy: None as far as Portugal is concerned. Material: 118 teeth. Localities: Santa Margarida (33), Esbarrondadoiro (77) and Vale de Zebro (8). Description: In this form, teeth are tiny. In occlusal view, the crown has a lozenge to an elliptical outline. The pitted labial surface is rounded near the transverse crest which appears in relief. The marginal angles are quite distinct. In the anterior teeth, the crown’s labial area presents a slight, ornamented depression. The lower edge from the labial visor is narrow, convex, and presents a slight protuberance at its midline. The lingual surface profile is concave. The median lingual ridge may be absent or is very thin and sharp. The con- cave crown visor overlaps the sulcus of the root. The root is two-lobed. The labio-lingually elongated lobes have an elliptical to triangular outline. The rather broad sulcus presents a nearly central foramen. The labial surface of the root is high and oblique in profile view. The lingual surface is reduced; its profile is concave. Fig. 2. 1, 2. Dasyatis gr. gigas (ESB; 15 a; U.E.). 1, lateral tooth: a, labial view; labial view; c, basal view. 3–5. Dasyatis gr. centroura (ESB; 15 b; U.E.). 3, anter labial view; b, basal view. Fig. 2. 1, 2. Dasyatis gr. gigas (ESB ; 15 a ; U.E.). 1, dent latérale : a, vue labiale b, vue labiale ; c, vue basilaire. 3–5. Dasyatis gr. centroura (ESB ; 15 b ; U.E.). 3, de latérale : a, vue labiale ; b, vue basilaire. only a little. In male teeth, the transverse crest is lingually developed into a long and pointed cusp. The labial surface, which presents a small longitudinal depression, is ornamented from the base to near the cusp’s apex. In profile view the labial surface is convex and the cusp tilts lingually. Marginal angles are quite distinct. The labial visor is strongly convex in its cen- tral part. The lingual surface is higher than in other species of the same genus and shows no ornamentation. The lingual visor is convex close by the root’s lobes, and presents a slight lateral depression near the marginal angles. The surface of the root is convex (and not flat as in other spp. from the same genus) and follows in smooth continuity the labial surface of the root, whose lateral profile is high and oblique. The sulcus presents a foramen in labial position. The lingual surface of the root is rather short. Discussion: This form clearly differs from all the hitherto described fossil Dasyatis species. It seems quite similar to the living D. marmorata and D. pastinaca, whose teeth are closely similar if not identical: despite our detailed observations on modern jaws it was not possible to find differences that could allow the distinction of the species concerned. It therefore is not possible to ascribe the fossil teeth to any one of these two species. That justifies its classification as above. Dasyatis cf. margaritella Compagno and Roberts, 1984 Fig. 4(2–5) Synonymy: None as far as Portugal is concerned. Material: 26 teeth. Localities: Santa Margarida (15), Esbarrondadoiro (9) and Vale de Zebro (2). Description: Teeth are small. The labial surface of the crown presents a reticulate ornamentation. The transverse crest bulges. Some specimens present distinct marginal angles. The convex labial visor has a central protuberance. The labial area shows a small lateral depression near the marginal angles. The lower edge of the labial visor is broad and convex. The lingual portion of the crown is high and has a concave lateral profile. The narrow median lingual ridge may be absent. The convex lingual visor largely overlaps the lobes of the root. The root lobes are low and incline lingually. The triangular- profiled lobes are elongated in labial-lingual orientation. The labial surface is wide and is slightly concave. The basal sur- faces of the root lobes are nearly flat, the lingual lobe being much reduced. The sulcus is deep and presents one foramen in a labial position. rofile; c, occlusal view; d, basal view. 2, anterior tooth of a male: a, profile; b, ooth: a, labial view; b, basal view. 4, lateral tooth: labial view. 5, lateral tooth: a, , profil ; c, vue occlusale ; d, vue basilaire. 2, dent antérieure de mâle : a, profil ; nt antérieure : a, vue labiale ; b, vue basilaire. 4, dent latéral : vue labiale. 5, dent A.C. Balbino, M.T. Antunes / Geobios 39 (2006) 747–755752 The teeth ascribed to males present a transverse crest that is more developed in its central part. The shorter, lingual surface shows no middle-lingual crest. In profile view, the crown has a convex labial surface and a concave lingual one. 3. Conclusions The dasyatid distribution in the Esbarrondadoiro Formation is shown (Table 1). w. 2 ral t aire A.C. Balbino, M.T. Antunes / Geobios 39 (2006) 747–755 753 Discussion: The above described teeth are similar to those of the living species D. margaritella observed by us. However, well-preserved specimens are scarce. Furthermore, species very close to D. margaritella are not yet described. Hence the taxo- nomic status of the fossil teeth seems quite problematical and justifies their classification as D. cf. margaritella. Dasyatis sp. Figs. 4(6–7) and 5(1, 2) Synonymy: None as far as Portugal is concerned. Material: 472 teeth. Localities: Santa Margarida (9), Esbarrondadoiro (456) and Vale de Zebro (7). Description: Teeth are large, and can be recognized by the very bulgy and strongly ornamented crown. The transverse crest is broad and pitted. Some teeth show sort of labial- lingually directed ribs. The labial visor of the crown is convex with a central prominence; its lower edge is broad and clearly separated from the root, without distinct marginal angles. The lingual surface of the crown is high and presents a broad, med- ian lingual ridge. The profile of the marginal–lingual surfaces is concave. The convex, lingual visor overlaps the root sulcus. The root lobes are separated by a broad and deep sulcus with two foramina in its central part. In profile view, the labial sur- face of the root is high and oblique, the lingual surface being rather short. In the lateral teeth the crown is mesio-distally elongated and ornamented. The lingual surface is shorter, and the convex lin- gual visor does not overlap the root, nor nearly so. The root lobes are somewhat broader, and the lingual surface of the root is more developed than the labial one. The teeth ascribed to males present a strong cusp. The labial surface is strongly ornamented, and the marginal angles are distinct. The edges of the labial and lingual visors are convex. The root is high, with thick and rather separated lobes. The sulcus presents a foramen in a labial position. Discussion: The fossil teeth have been compared to those from the living species D. pastinaca, D. marmorata, D. margaritella, D. violacea, D. centroura and D. gigas. How- ever, the fossil teeth are very different from all these species by their robustness, by the crown morphology and by their orna- mentation. The fossil teeth may represent a new species, but a better knowledge of the dentition of living species is needed before any decision can be taken. It therefore seems better to leave this species in open nomenclature. Fig. 3. 1, 2. Dasyatis gr. centroura (ESB; 15 b; U.E.). 1, lateral tooth: basal vie pastinaca (ESB; 15 c; U.E.). 3, lateral tooth: a, labial view; b, basal view. 4, late 6, anterior tooth: a, labial view; b, profile; c, occlusal view. Fig. 3. 1, 2. Dasyatis gr. centroura (ESB ; 15 b ; U.E.). 1, dent latérale : vue basil pastinaca (ESB; 15 c; U.E.). 3, dent latérale : a, vue labiale ; b, vue basilaire. 4, dent basilaire. 6, dent antérieure : a, vue labiale ; b, profil ; c, vue occlusale. In the selachian ichthyofauna of the Latest Miocene of the Alvalade Basin (Esbarrondadoiro Formation), myliobatiform remnants represent about 34% of the whole (Antunes et al., 1999). Among the myliobatiforms, Dasyatidae are abundant, with about 46%. The Miocene deposits of the Alvalade Basin are particularly rich in modern type, close to still extant forms (Balbino, 1995; Antunes et al., 1999). This is confirmed by the presence of different species of Dasyatis. Effectively, Dasyatis gr. cen- troura, D. marmorata and D. pastinaca live in the Mediterra- nean and the Atlantic, while D. gr. gigas and D. margaritella are North-Eastern Atlantic dwellers. From the thermic point of view there are important data. The genus Hemipristis is entirely absent from the Alvalade Basin, where there are almost no tiger sharks (Galeocerdo) remnants. However, these warmth-loving forms still occur in the earlier, Tortonian associations from Lisbon, although Hemipiristis is quite rare, Galeocerdo being rather scarce (Bal- bino, 1995; Antunes and Jonet, 1970). This points out to a decrease in the prevailing thermic conditions, which seems cor- roborated by the myliobatiforms. As it has been shown, the Dasyatis species are still abundant and varied among the Alva- lade basin fauna in contrast to the rarity of Raja, a genus repre- sented but in a small percentage (10%) (Balbino, 1995). Raja otherwise suggests temperate to cold (or deeper and hence rather cold) waters, and this is precisely the opposite of the current, prevailing thermic situation. In summary, moderately warm water conditions predominated, less than in Tortonian of Lisbon, more than what exist today. Another meaningful point results from the quantitative representation of the dasyatid taxa among the three studied localities. As it can acknowledged (Table 1), some forms (Dasyatis gr. centroura, D. marmorata / pastinaca and D. cf. margaritella) occur in nearly identical or in not meaningfully different percentages among the localities under study. How- ever, there are strong differences as far as Dasyatis gr. gigas and D. sp. are concerned: the Esbarrondadoiro pattern is very different from those from Santa Margarida and Vale de Zebro, which are nearly identical between them. D. gr. gigas may sug- gest the occurrence of less warm waters, and this is very much compatible with other data as hexanchid and squaliform occur- rences that indicate a deeper environment and hence colder waters for Esbarrondadoiro. The Alvalade basin fossil forms of Dasyatis had a litoral/ neritic bathymetric distribution and a benthic way of life as shown by comparison with close-by living ones Figs. 5. , anterior tooth of a male: a, labial view; b, profile. 3–6. Dasyatis marmorata / ooth: a, labial view; b, basal view. 5, lateral tooth: a, labial view; b, basal view. . 2, dent antérieure de mâle : a, vue labiale ; b, profil. 3–6. Dasyatis marmorata / latérale : a, vue labiale ; b, vue basilaire. 5, dent latérale : a, vue labiale ; b, vue A.C. Balbino, M.T. Antunes / Geobios 39 (2006) 747–755754 , south alade, S ida ecimens e: a, lab : a, lab ; b, bas mâle : ire. 3, d dent lat Table 1 Latest Miocene Dasyatids from the Esbarrondadoiro Formation, Alvalade Basin Dasyatidés de la Formation Esbarrondadoiro (Miocène terminal), Bassin d’Alv Taxa Localities Santa Margar Warm (w) and temperate (t) waters Number of sp and % Dasyatis gr. gigas w > t 118/45.2 Dasyatis gr. centroura w 96/36.8 Dasyatis marmorata/pastinaca w 23/8.8 Dasyatis cf. margaritella w 15/5.7 Dasyatis sp. – 9/3.4 Totals 5 261/99.9 Fig. 4. 1. Dasyatis marmorata/pastinaca (ESB; 15 c; U.E.); lateral tooth of a mal (ESB; 15 d; U.E.). 2, anterior tooth: a, labial view; b, basal view. 3, lateral tooth the crown. 6, 7. Dasyatis sp. (ESB; 15 e; U.E.). 6, lateral tooth: a, labial view Fig. 4. 1. Dasyatis marmorata / pastinaca (ESB ; 15 c ; U.E.) ; dent latérale de margaritella (ESB ; 15 d ; U.E.). 2, dent antérieure : a, vue labiale ; b, vue basila dent latérale : détail de la couronne. 6, 7. Dasyatis sp. (ESB ; 15 e ; U.E.). 6, Acknowledgements This work results from research concerning the Line no. 1 of the Centro de Estudo Geológicos of the Universidade Nova de Lisboa. References Antunes, M.T., Balbino, A.C., Cappetta, H., 1999. Sélaciens du miocène term- inal du bassin d’Alvalade (Portugal). Essai de synthèse. Ciências da Terra (UNL). Lisboa 13, 115–129. Antunes, M.T., Jonet, S., 1970. Requins de l’Helvétien supérieur du Tortonien de Lisbonne. Revista da Faculdade de Ciências da Universidade de Lisboa 2, C 16 (1), 119–280. Antunes, M.T., Mein, P., 1995. Nouvelles données sur les petits mammifères du miocène terminal du bassin de Alvalade, Portugal. Comunicações do Instituto Geológico e Mineiro 81, 85–96. Balbino, A.C., 1995. Seláceos (Pisces) do Miocénico terminal da Bacia de Alvalade (Portugal)/Sistemática, Ecologia, Paleoambientes, Comparação com faunas actuais. Dissertação de Doutoramento. Universidade de Évora. Bigelow, H.B., Schroeder, W.C., 1953. Fishes of the western north Atlantic, part 2, Mem. Sears Foundation for Marine Research 1 (2), 1–588. Ca Ca Ca Co Jon Jon Nis Pro Ser Fig. 5. 1, 2. Dasyatis sp. (ESB; 15 e; U.E.). 1, lateral tooth: a, labial view; b, basal v Fig. 5. 1, 2. Dasyatis sp. (ESB ; 15 e ; U.E.). 1, dent latérale : a, vue labiale ; b, vue ern Portugal ud du Portugal Esbarrondadoiro Vale de Zebro Σ % Number of specimens and % Number of specimens and % 260/21.9 62/47.7 440 27.88 385/32.4 51/39.2 532 33.71 77/6.5 8/6.2 108 6.84 9/0.8 2/1.5 26 1.65 456/38.4 7/5.4 472 29.91 1187/100 130/100 1578 99.99 ial view; b, basal view; c, detail of the crown. 2–5. Dasyatis cf. margaritella ial view; b, basal view. 4, lateral tooth: labial view. 5, lateral tooth: detail of al view. 7, lateral tooth: labial view. a, vue labiale ; b, vue basilaire ; c, détail de la couronne. 2–5. Dasyatis cf. ent latérale : a, vue labiale ; b, vue basilaire. 4, dent latérale : vue labiale. 5, érale : a, vue labiale ; b, vue basilaire. 7, dent latérale : vue labiale. A.C. Balbino, M.T. Antunes / Geobios 39 (2006) 747–755 755 ppetta, H.C., 1970. Les Sélaciens du Miocène de la région de Montpellier. Palaeovertebrata, Mémoire extraordinaire, 1–139. ppetta, H.C., 1980. Modification du statut générique de quelques espèces de sélaciens crétacés et tertiaires. Palaeovertebrata 10, 29–42. ppetta, H.C., 1987. Mesozoic and Cenozoic Elasmobranchii Chondrichthyes II. Gustav Fischer Verlag, Stuttgart. mpagno, L.J.V., Roberts, T.R., 1984. Marine and freshwater stingrays (Dasyatidae) of West Africa, with description of a new species. Proceed- ings of the California Academy of Sciences 43, 283–300. et, S., 1968. Notes d’ichthyologie Miocène portugaise V – Quelques Batoïdes. Revista da Faculdade de Ciências da Universidade de Lisboa 2, C 15 (2), 233–257. et, S., 1978. Le Tortonien supérieur (T VII-b) des environs de Fonte da Telha (Péninsule de Setubal) et ses faunes. Comunicações dos Seviços Geológicos de Portugal. Lisboa 63, 13–51. hida, K., 1990. Phylogeny of the suborder Myliobatidoidei. Hokaido Uni- versity 3, 1–108. bst, J., 1877. Beiträge zur Kenntniss der fossilen Fische aus der molasses von Baltringen. II: Batoïdei A. Günther. Jahreshefte des Vereins für vater- ländische Naturkunde in Württemberg 33, 69–103. ralheiro, A.M.R., 1954. Contribuição para o conhecimento da fauna ictio- lógica do Miocénico marinho de Portugal Continental. Revista da Facul- dade de Ciências da Universidade de Lisboa 2, C 4 (1), 39–119. iew. 2, lateral tooth of a male: occlusal view. basilaire. 2, dent latérale de mâle : vue occlusale. Latest Miocene Dasyatidae (Neoselachii, Batomorphii) from the Alvalade Basin, Portugal Introduction Systematics Conclusions Acknowledgements References