New material of Diacodexis (Mammalia,Artiodactyla) from the early Eocene of Southern Europe

Diacodexeidae are the first representatives of Artiodactyla in the fossil record. Their first occurrence is at the very base of the Ypresian (earliest Eocene, 56.0 Ma) with Diacodexis, a genus well diversified during the early Eocene in Europe, especially during the MP7–MP8 + 9 interval. However, most of European species are documented by scarce material, retrieved from single localities. In this work, we describe new Diacodexis material from ～MP7 and ～MP8 + 9 localities of Southern Europe, including material of D. antunesi from Silveirinha, considered as the most primitive European Diacodexis species, and material from three localities from Southern France (Fordones, Palette, and La Borie). The new material documents Diacodexis premolar morphology and deciduous dentition which bear potentially important phylogenetic information, as well as astragali, including a specimen from Silveirinha that constitutes the earliest occurrence of an astragalus of the genus Diacodexis in the European fossil record. Investigation of the enamel microstructure reveals that early European species had a simple enamel pattern with one-layered Schmelzmuster composed of ‘basic’ radial enamel only, instead of the two-layered Schmelzmuster (thin radial enamel + thick layer of Hunter-Schreger bands) observed on North American species and so far considered to represent the primitive condition within Artiodactyla. In accordance with previous studies, our observations highlight that Diacodexis gigasei from Belgium is morphologically closer to the North American species D. ilicis than to D. antunesi from Portugal. The latter species, together with D. aff. antunesi from Fordones, appears to be morphologically closer to the Asiatic taxa D. indicus and D. pakistanensis. Finally, we found numerous similarities between D. cf. gigasei from Palette and D. gigasei, a result that challenges the intra-European provincialism that characterizes the earliest Ypresian. Diacodexis gigasei could be one of the rare species shared by the northwestern and southwestern European bioprovinces.     

A new species of Dyrosaurus (Crocodylomorpha, Dyrosauridae) from the early Eocene of Morocco: phylogenetic implications

A new dyrosaurid is described from the Ypresian of the phosphatic deposits of the Oulad Abdoun Basin of Morocco. It is based on numerous cranial and postcranial remains, allowing an almost complete reconstruction. This new Dyrosaurus species, Dyrosaurus maghribensis sp. nov. , is currently only known from Morocco. It differs from D. phosphaticus , present in contemporaneous levels of Algeria and Tunisia, by several autapomorpies, including a smooth dorsal margin of the parietal and widely opened choanae. A phylogenetic analysis, using 47 taxa and 234 morphological characters, shows the dyrosaurids as the sister taxon of pholidosaurids, which include Elosuchus , Sarcosuchus , Terminonaris and Pholidosaurus , and the thalattosuchians. Goniopholididae is a non-monophyletic group; however, if dyrosaurids are not included in the analysis, the result differs and the goniopholidids form a distinct clade. If Thalattosuchia is excluded, both Goniopholididae and Pholidosauridae become paraphyletic assemblages. Thus, phylogenetic problems remain with respect to longirostrine clade, and more attention should be paid to resolving their evolutionary relationships amongst the crocodyliforms.  © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society , 2006, 148 , 603–656.     

A new basal tapiromorph (Perissodactyla, Mammalia) from the middle Eocene of Myanmar

A new genus and species of tapiromorph, Skopaiolophus burmese nov. gen., nov. sp., is described from the middle Eocene Pondaung Formation in central Myanmar. This small form displays a striking selenolophodont morphology associated with a mixture of primitive “condylarthran” dental characters and derived tapiromorph features. Skopaiolophus is here tentatively referred to a group of Asian tapiromorphs unknown so far. The occurrence of such a form in Pondaung suggests that primitive tapiromorphs might have persisted in southeast Asia until the late middle Eocene while they became extinct elsewhere in both Eurasia and North America.     

A new species of Neochelys (Chelonii,Podocnemididae) from the Ypresian (Early Eocene) of the South of France

The known European record of the Gondwanan group Podocnemididae begins in the Early Eocene. Neochelys underwent a rapid diversification and was an abundant and diverse representative of this group during the Eocene of Europe from the Early Ypresian to at least the Bartonian. However, several of its species are poorly known, and the phylogenetic relationships among them are poorly understood. A complete shell from the Ypresian of Hérault (southern France) is described here. It is assigned to a new species, Neochelys liriae. The availability of characters of N. liriae is tested by comparison with the other European species, which appear to be well differentiated, and particularly with the neighbouring French taxa N. eocaenica and N. laurenti, of which the latter has been little known to this point.     

Systematics and diversity of the giant soft-shelled turtles (Cryptodira,Trionychidae) from the earliest Eocene of Belgium

In 1909, the famous paleontologist Louis Dollo identified two putative new species of giant soft-shelled turtles from the lowest Eocene record of Belgium, ‘Trionyx erquelinnensis’ and ‘Trionyx levalensis’, from Erquelinnes and Leval, respectively. However, these proposals did not meet the requirements of the International Code of Zoological Nomenclature, so they were considered as nomina nuda. The information on these specimens or about any other specimen of this lineage of giant turtles from the Belgian record is currently extremely limited. Relatively scarce material from giant trionychids has been described for the lower Eocene record of other European regions. Considering the available information, all the European material has recently been recognized as belonging to the genus Axestemys, which has a North American origin, and possibly attributable to a single species, Axestemys vittata, which currently lacks a diagnosis. Numerous and well-preserved Belgian specimens are deposited in the Royal Belgian Institute of Natural Sciences. In addition to the cited individuals from Erquelinnes and Leval, additional specimens from both localities, as well as others from Orp-le-Grand, are part of this collection. These specimens, found between the decades of 1910 and 1930, have been recently restored, and their study is carried out here. The presence of Axestemys vittata in Belgium (in Leval and Orp-le-Grand) is confirmed. Knowledge about this species is significantly improved, and a diagnosis is proposed. However, the hypothesis proposed by Dollo is here confirmed, this species being not the only identified in the Belgian record. So, Axestemys erquelinnensis nov. sp. is defined based on the carapace from Erquelinnes known by Dollo, suggesting that the genus probably reached Europe during the Paleocene Eocene Thermal Maximum.     

The paroxyclaenidae (Mammalia) and a new form from the early Eocene of Palette,France

A new genus and species of the Paroxyclaenidae is described,Merialus martinae, and the family is reviewed and divided into two subfamilies, the Paroxyclaeninae and the Merialinae, new. The two species ofDyspterna are attributed to two genera, of whichEuhookeria is new; Dyspterninae Kretzoi, 1943, is resurrected to includeDyspterna woodi, Cryptopithecus andKochictis; the subfamily is referred to the Pantolestidae.Vulpavoides is considered valid and includes“Russellites” simplicidens;V. vanvaleni n. sp. is described from Bouxwiller.Kiinkerishella andDulcidon are discussed, as are possible relationships between the Ptolemaiidae and the Merialine paroxyclaenids.     

A primitive ceratomorph (Perissodactyla, Mammalia) from the early Tertiary of Europe

Hyracotherium cuniculus Owen, 1842, from Palaeocene–Eocene boundary strata in NW Europe, was formerly considered to be an equoid. It is here placed in Cymbalophus gen. nov. , in the perissodactyl suborder Ceralomorpha, which comprises tapiroids and rhinocerotoids, as one of its most primitive and earliest members. This revised classification is based on an evaluation of its advanced versus primitive character states, and comparison with a variety of other primitive perissodactyls. Cheek tooth occlusal relationships in the early stages of ceratomorph evolution are discussed and contrasted with the fundamental perissodactyl morphotype as represented in Hyracotherium. It is suggested that the Lophiodontidae (formerly considered to be ceratomorphs; may be more closely related to chalicotheres; that the Isectolophidae may be the sister-group of chalicotheres and lophiodonts plus ceratomorplis; and that Cymbalophus is the most primitive known member of the newly restricted Ceratomorpha. Cymbalophus is placed tentatively in the family Helaletidae.     

The oldest mammals from Antarctica,early Eocene of the La Meseta Formation,Seymour Island

New fossil mammals found at the base of Acantilados II Allomember of the La Meseta Formation, from the early Eocene (Ypresian) of Seymour Island, represent the oldest evidence of this group in Antarctica. Two specimens are here described; the first belongs to a talonid portion of a lower right molar assigned to the sparnotheriodontid litoptern Notiolofos sp. cf. N. arquinotiensis. Sparnotheriodontid were medium‐ to large‐sized ungulates, with a wide distribution in the Eocene of South America and Antarctica. The second specimen is an intermediate phalanx referred to an indeterminate Eutheria, probably a South American native ungulate. These Antarctic findings in sediments of 55.3 Ma query the minimum age needed for terrestrial mammals to spread from South America to Antarctica, which should have occurred before the final break‐up of Gondwana. This event involves the disappearance of the land bridge formed by the Weddellian Isthmus, which connected West Antarctica and southern South America from the Late Cretaceous until sometime in the earliest Palaeogene.     

The phylogeny of the Rhinocerotoidea (Mammalia, Perissodactyla)

A new phylogeny of the Superfamily Rhinocerotoidea is proposed, based upon an analysis of shared derived characters of the skull, teeth and skeleton. Hyrachyus is considered the primitive sister-taxon of the three rhinocerotoid families (Amynodontidae, Hyracodontidae, Rhinocerotidae), and the amynodonts appear to be the sister-group of hyracodonts and rhinocerotids. The relationships of primitive hyracodonts and rhinocerotids are clarified. Menoceras and Pleuroceras (new Subfamily Menoceratinae) are removed from the Diceratheriinae, since they appear to be more closely related to higher rhinoceroses than they are to Diceratherium. Of the three major monophyletic groups of higher rhinocerotids (aceratherines, teleoceratines and rhinocerotines), the last two groups are more closely related to each other than either is to aceratherines. Toxotherium and Schizotheroides are tentatively removed from the Rhinocerotoidea and placed in the Lophiodontidae.     

Variation in the shape of the palate in adapis (Eocene,Adapiformes) compared with living primates

This study is a quantitative analysis of the variation of palate shape inAdapis, an Eocene fossil primate. The observed coefficients of variation of palate dimensions inAdapis are statistically greater than in most living primate species. The histogram of the index (palate width/palate length) emphasizes the great heterogeneity in the fossil species. Two allometric scalings of palate dimensions (length and width) on condylobasal length were performed. Plots of perpendicular distances from the two regression lines show that living species are always more concentrated around the middle of the diagrams than isAdapis. The space occupied byAdapis is as great as, or even greater, than those occupied by living genera. Hence, several species are certainly presenent in the fossil sample, and the varieties distinguished by Stehlin in 1912, are in fact valid species names.     

New remains of Egatochoerus jaegeri (Mammalia,Suoidea) from the late Eocene of Peninsular Thailand

Abstract: We report here additional remains referred to Egatochoerus jaegeri from the late Eocene locality of Ban Mark in the Krabi basin, Thailand. The new material described, comprising upper and lower cheek teeth, deciduous premolars and partial cranial remains, makes E. jaegeri the best‐documented Eocene representative of Old World Suoidea at present. Detailed study and comparison of their cheek teeth structure reveal a homogeneity of the molar crest and groove patterns of Old World and New World Palaeogene suoids.     

Additional vertebral material of Thaumastophis (Serpentes: Caenophidia) from the early Eocene of India provides new insights on the early diversification of colubroidean snakes

The Ypresian Cambay Shale Formation at Vastan, Mangrol, and Tadkeshwar lignite mines in Gujarat, western India, has yielded a rich vertebrate fauna including madtsoiid, palaeophiid, booid, and colubroidean-like snakes. The latter are particularly abundant, but their systematic affinities are difficult to resolve. Here we describe new specimens of the colubroidean-like snake Thaumastophis missiaeni, including anterior, middle, and posterior trunk vertebrae, as well as caudal vertebrae. The combination of primitive and derived caenophidian and colubroidean vertebral characters confirms Thaumastophis as the earliest known stem-colubriform snake while Procerophis, from the same beds, is more derived and considered to represent a crown-Colubriformes. Additionally, Thaumastophis shares with Renenutet enmerwer from the late Eocene of Egypt a unique combination of vertebral characters that suggests an exchange with North Africa was possible along the southern margin of the Neotethys. We erect the new family Thaumastophiidae for Thaumastophis and Renenutet on the basis of their shared derived vertebral morphology.     

The first record of Sirenia (Mammalia) from the early Oligocene of the Paratethys

Abstract

Sirenians have been extensively recorded from the Mediterranean and west European localities but there are only few finds to the east of it, from the area covered by the Paratethys. For the early Oligocene, to our knowledge, there are no published records of sirenians from inner seas of the Old World. Here we report a specimen of Dugongidae indet., consisting of two partial vertebrae and 12 fragments of ribs, collected in a manganese ore mine in Ukraine and dated as the earliest Oligocene (33–32 Ma). The specimens, as preserved, did not differ in morphology and size from ‘Halitherium schinzii’ and therefore can belong to Kaupitherium, at present the single early Oligocene genus recorded from Europe. However, its vertebral and rib anatomy is not specific for Kaupitherium, so we identify it only by family level. The marks of scavenging on a rib possibly are due to gastropod or bivalve mollusks. The sea, as suggested from biotic data, had a temperate or subtropical climate, relatively cold waters and high diversity of pelagic and deep-water habitats. Thus, the onset of the Oligocene was a period when sirenians could enter temperate inner Eurasian waters, a marginal area in their worldwide dispersal.