The mammalian faunas of the Paleogene of Europe and their localities are reviewed with comments on problems of European stratigraphy (epoch, stage and substage limits) and on the possibilities of faunal migrations. Radiometric dating is discussed. A stratigraphic scale for the Paleogene is presented, as well as a refined system of sequential faunal levels. 

A new species, Otonycteris rummeli nov. sp., is described from the Late Miocene site Gritsev (MN 9) in the Ukraine. Otonycteris rummeli nov. sp. differs from those of most vespertilionids, except recent Otonycteris, Antrozous and Early Miocene Karstala silva, in having a well-developed entocingulid at the foot of the trigonid valley in the lower molars. The morphological resemblance of Otonycteris, Antrozous and Karstala is apparently a case of convergence in the evolution of the Old and New Worlds bat faunas. From at least the Middle Miocene the range of Otonycteris distribution spread to the whole of Central Europe and such a situation continued during the whole Late Miocene. This indicates a more arid climate in Europe during the Upper Miocene compared to the Quaternary. The reduction of the distribution range of Otonycteris and its extinction in most of the territory of Europe could have been caused by the global climatic cooling and increasing glacial cycle amplitude during the onset of the Quaternary. 

The dentition of selachians is characterized by an often very pronounced heterodonty involving a great morphological diversity. Despite this fact, the dentitions of selachians can be grouped in a rather reduced number of dental types corresponding to trophic adaptations: grasping, tearing, cutting, crushing, grinding and grasping-grinding type. The numerous exemples of convergence and parallelism that can be observed in fossil selachians and between Recent and fossil ones is the result of this reduced number of dental types. These dental specialisations allow to try a reconstruction of the way of life of fossil forms. 

Some new material of percrocutas from the late Miocene of Axios valley (Macedonia, Greece) is studied. They have been found in the locality of "Pentalophos 1" (PNT). The material has been described and compared with the known late Miocene percrocutas of Eurasia. This comparison indicates that it can be identified as Dinocrocuta gigantea (SCHLOSSER, 1903). A maxilla of a percrocuta, named ”Hyaena" salonicae, was found in the same area (Andrews, 1918). "Hyaena" salonicae is smaller than the PNT material. It is also compared with other material from Eurasia while its taxonomic and age problems are discussed. It belongs to Dinocrocuta and shows close relationships with D. robusta and D. senyureki; its age can be considered as late Vallesian-early Turolian. The age of the locality PNT is also discussed and a possible Vallesian age is proposed for it. 

The anatomy of Norselaspis glacialis n.g., n.sp., a primitive kiaeraspidian from the Lower Devonian of Spitsbergen, is described on the basis of spécimens studied by grinding sections or prepared with dilute formic acid. This study yielded some new anatomical details, including the presence of a canal prolonging posteromedially the canal alloted to the facial nerve by Stensiö. This posterior prolongation of the « facial canal ›› into the posterolateral part of the labyrinth cavity is consistent with the hypothesis put forward by Allis, Lindström, Jefferies and Whiting, that this canal housed the glossopharyngeus nerve. Furthermore, in N. glacialis, the foramen usually referred to as the foramen for the œsophagus opens posteriorly into a cavity in the postbranchial wall, referred to here as the intramural cavity, and which is interpreted as having housed the heart. Consequently, the œsophagus probably accompanied the dorsal aorta through the aortic canal. Finally, the foramen generally interpreted as having transmitted the ventral afferent arterial trunk is here considered as having housed the hepatic vein, which emptied into the venous sinus of the heart. The ventral afferent arterial trunk may thus have passed through the former «œsophageal ›› foramen.
The problem of the position of the dorsal nerves in the Osteostraci is discussed, and it is suggested that the three foremost nerve canals opening into the oralobranchial cavity housed the maxillary ramus of the trigeminus, the facial nerve and the glossopharyngeus nerve respectively. The mandibular ramus of the trigeminus must have accompanied one of the two foremost nerves, but for the moment it is impossible to decide which.
The problem of the nature of the interbranchial crests of the Osteostraci is briefly discussed. Comparison with the branchial apparatus of the Petromyzontida does not support the hypothesis that the interbranchial crests are part of the branchial arches, incorporated into the endoskeletal shield. A different hypothesis is proposed, that the branchial skeleton of the Osteostraci was situated entirely inside the oralobranchial cavity, and was attached to the endoskeletal shield only by the ventromedial processes. The grooves classically allotted to the efferent branchial arteries would thus have housed extrabranchial arteries, branching off from the dorsal aorta, and irrigating the ventral branchial musculature.
A phylogeny and a classification of the kiaeraspidians are proposed. The evolution of this monophyletic group is characterized by, e.g., reduction of cornual processes, shortening of the abdominal division of the shield, subdivision of the lateral fields, and enlargement of the supraoral fossae.
The phylogenetic position of the kiaeraspidians within the Osteostraci remains uncertain. Their sister-group may be either the benneviaspidiens or the thyestidians, or Thyestes alone (in which case they would have to be included within the thyestidians). 

The genus Plagiolophus is documented, almost solely in Western Europe, from the middle Eocene up to the mid Oligocene (MP 12 to MP 25), i.e. more than for 15 MY. Seventeen species are now recorded whose two of them are new, P. ringeadei nov. sp. and P. mamertensis nov. sp. Some anatomical variations and the deflection of certain evolutionary trends justify the distinction of three subgenera, Paloplotherium, Fraasiolophus nov. and Plagiolophus s.s. The genus displays a wide range in size and weight (between 10 and 150 kg). The detailed description of the skull of several species is here given for the first time.
Despite important evolutionary drifts during this long time span, the dentition shows a great structural homogeneity, which renders difficult the determination of fragmentary specimens or isolated teeth. It is characterized by a great heterodonty; premolars are little molarized and present a certain regression through time with paradoxically some progress in the molarization. The hypsodonty increases: the first Plagiolophus are hardly less brachyodont than Propalaeotherium, and the last ones are nearly as hypsodont as Merychippus from the early Miocene. The upper molars change from a wide crown pattern, with an open occlusal surface, lightly oblique transverse lophs and rounded internal cusps, to a narrower pattern, with a frontally constricted occlusal surface and internal lophs aligned parallel to the ectoloph. The M3/3 become always longer.

The dental enamel displays horizontal Schreger-bands with imprecise limits occupying only the middle part of the enamel layer. The dentine is remarkable by its high rate of pericanalicular dentine. The crown cementum, lacking in earlier forms, increases to the point where it fills the occlusal valleys of the
teeth.

The masticatory musculature shows a increasing prominence of the temporal, with probably an important role devoted to the pterygoid muscles in lateral movements related to a two-phase type of chewing.

The evolution of the dentition, of the masticatory musculature and of the repartition of masticatory forces indicate that the Plagiolophus have known different diets through their long evolutionary history; at first browsers they became mixed feeders and finally grazers. Their relatively long neck allowed these animals to reach different vegetal layers. The strength of the nuchal crests also suggests that they were able to have strong backwards movements of the head to pull up their food.

This evolution of diet seems related to the slow degradation of environmental conditions attested during this period in western Europe, with the generalization of more open landscapes, increasing aridity and more marked seasons.

Besides, a remodeling of the face is ontogenetically and along time observed, in relation with the evolution of the masticatory apparatus and especially with that of the mandibular lever arm. The postcanine diastemata become longer in the course of evolution; the free extremities of the nasals are always relatively long which contradicts the hypothesis according to which Paloplotherium may have had a trunk. At last the lineage Fraasiolophus can be distinguished by the presence of a deep malar fossa, probably related to a strong development of the maxillo-labialis superior muscle.

The orbit is always large and tends to increase in size, which indicates a good development of the vision and its increasing role in the life relations. A peculiar type of epitympanic sinus could have been used as a resonance chamber insuring a certain amplification of sounds before their transmission to the eardrum. The endocranial cast reveals a relatively large brain with an advanced degree of gyrencephaly. Beside the role eventually played in food research and social relations, these neurophysiological abilities, also related to an advance in cursorial fitness, could have contributed to the survival of these animals facing the predation pressure of the first fissipede carnivores and the competition with new immigrant herbivores after the "Grande Coupure".
On the basis of some shared apomorphies with the Pachynolophinae, which prevent from considering the latter as Equidae (molarization of the premolars, reduction of the premaxilla dorsal apophysis, peculiar epitympanic sinus, splitting of the jugular process), the hypothesis of an autochthonous origin of Plagiolophus issued from a form near Propalaeotherium, is once again proposed and discussed. Finally, intra-generic relationships are taken into consideration. 

In recent years, the techniques of chemical preparing have permitted a rich paleontologic material to be obtained from the phosporitic sediment of Bouzigues (Hérault, France). The fauna of this locality is comprised of quite varied microvertebrates, amphibians, reptiles, birds, mammals. Twenty five species of the latter, belonging to seven orders, are today known from the site. Among them, the numerous rodents have allowed L. Thaler to chronologically situate this fauna in the Zone of Laugnac (<< late Aquitanian ›> of some authors).
The chiropterans are, with the rodents, the best represented of the locality's mammals. Three families comprise the bat fauna, with nearly complete dominance by one of them (Hippoxideridae) over the two others (Megadermatidae and Vespertílionidae)
Six forms are described, of which three are new species and one a new sub-genus.
Megaderma braillomi n. sp., an animal of rather large size, shows like the Miocene megaderms several evolved dental characters, translating the adaptation of these animals to a partially carnivorous regime. The Bouzigues species seems, however, to represent a particular lineage.
Hipposideros (Brachipposideros n. subgen.) dechaseauxi n. sp. and Hípposideros (Brachipposideros) cf. collongenris Depéret, small sized forms, belong to a group rather well represented in the late Oligocene and early Miocene of Europe, and not distinguished until now within the genus Hipposideros.
Hipposideros (Pseudorhinolophus) bouziguensis n. sp., is the most abundant mammal in the locality and, occuring at the Oligocene-Miocene limit, the last representative known of the subgenus Pseudorhínolophus, common in Europe from the middle Eocene.
However, beyond Neogene and Quaternary times, certain among the numerous living species of Hipposideros are close to Pseudorhinolophus and others to Brachipposíderos. 'This fact would in the future justify a global revision of the genus, on the basis of comparative anatomy of the squeleton and of the teeth.
The bat fauna of Bouzigues is completed by two small Vespertilionidae, rare forms, Myoris sp. I and sp. II.
  

Near "la Lieude", in the Lodève basin, more than a thousand of footprints are distributed in a twenty of trackways which amounts to 220 m length. They have been found on calcareous siltstone level in the B site named also "Réserve Naturelle Volontaire". This last is located in the Saxonian summit dated Upper Permian. "La Lieude" tracks are described by using statistical methods then they are compared with others from the world Permian. What allows to distinguish 4 following ichnotaxa: Lunaepes ollierorum nov.ichnosp., Merifontichnus thalerius nov. ichnogen. and nov. ichnosp., Planipes brachydactylus nov.ichnosp. and Brontopus circagiganteus nov. ichnosp. All these traces are attributed with possibility or probability to Therapsida or to Therosauria, except Brontopus circagiganteus nov. ichnosp. that could be due to Caseamorpha. All these animals whose sizes have been estimated between l and 5 m lived probably in a playa environment.The biological and sedimentological data from "la Lieude" footprints levels compared with informations provided by the tracks orientations, suggest the following scenario. Animals coming from the North have crossed a sandy channel bank with plants zones by directing to the South for the majority. Maybe, they were going to the lacustrine part of the playa, close to "la Lieude" footprints that they have just trampled on. 

Two new finds of pantodont materials from southern China, assigned to Archaeolambda, are described in this paper. One, a new species from the Nanxiong Basin, Guangdong Province, is similar to Alcidedorbignya inopinata from the early Paleocene of Tiupampa, Bolivia in size. It provides reliable evidence of the occurrence of Archaeolambda in the early-middle Paleocene of southern China. The second find includes specimens of Archaeolambda sp. cf. A. planicanina from the ?late Paleocene of Hengyang Basin, Hunan Province, which are the first record of a fossil mammal from the area near Hengyang city. The only vertebrate fossils previously found here were two genera of crocodiles discovered in 1938. This find sheds new light on the local biostratigraphy. 

Revision of ancient collections and study of new material from Provenchères-sur-Meuse (Rhaetian) lead to signíficant changes in the faunal list of this site. This bring to us important information about the effect of the rhaetian transgression on the evolution of the faunas at this period of time. However, study of
vertebrate microremains is always difficult and some points remain obscure, like the origin of the prosauropods which yield some remains at Provenchères-sur-Meuse and the survival in Europe during the Rhaetian of temnospondyl arnphibians different from the Plagiosauridae.

  

The discovery in the Turolian (Late Miocene) of Dytiko 3 (Macedonia, Greece) of a complete skull with mandibles and cervical vertebrae, atlas and epistropheus, is a very important contribution to the knowledge of the subfarnily Chalicotheríinae. After the description, the comparison with other specimens of Miocene chalicotheres permits the revival of the generic name Macrotherium with a new species M. macedonicum. This genus is mainly characterized by a short snout and an inflated cerebral skull. It coexists during the Miocene with Chalicotherium. A cladistic analysis leads to conclusion that the species which has been described from the Early Middle Miocene of Rusinga must be identified as the type-species of a new genus: Butleria.





  

In dilambdodont molars the primitive crest between paracone and metacone (centrocrista) is represented by a pair of crests that join the mesostyle (postparacrista, premetacrista). The cutting action of these crests against the crests of the hypoconid is described. Dilambdodonty is a derived adaptation for greater cutting efficiency. It has evolved several times and in more than one way. 

The Causses" is a near 3400 km2 large plateau located in the south of France. Here the first dinosaur footprints where found in 1935. After this, this area has yielded an ever-increasing number of ichnites now in excess of 500 specimens. These latter, 15 to 50 cm long, tridactyl or tetradactyl footprints of generally biped animals, were discovered at the surface of Hettangian to lower Sinemurian dolomite layers within 4 distinct stratigraphic units. The 35 sites bearing ichnites are located on the plateau margin. For the first time, morphologic characters studied through descriptive statistic methods with the usual parameters and classical Student and Snédecor tests, allowed us, to divide the whole set of biped traces into 6 ichnospecies. Their definitions are further constrained by multivariate statistical results using Principal Component Analysis (PCA), Factor Analysis of correspondances (FAC) and Discriminant Analysis (DA). All have confirmed the morphologic observations. So that now, the following taxa are identified : Grallator variabilis, G. lescurei, G. sauclierensis, G. minusculus, Eubrontes giganteus, Dilophosauripus williamsi, cf. Moraesichnium, Orníthopus fabrei nov ichnosp. The more immediately visible differences relate to the interdigital II-IV divarication and the digit length ratio. To this panel, we must add Batrachopus deweyi and shapes suggesting Trisauropodichnus and/or Anomoepus. Among all ichnite associations described in the lower Liasic, the New England assemblage presents the most affinities with ours. It shows the ichnotaxa Grallator, Dilophosauripus, Eubrontes, Batrachopus without forgetting Ornithopus fabrei nov. ichnosp. which is close to Ornithopus gallinaceus from the Massachusetts and Connecticut basins. On comparing the present early Jurassic ichnofauna of the Causses with the ones of the Middle and Upper Triassic formations of the eastem border of the Massif Central (France), it appears that tridactyl footprints become more and more numerous and large from Triassic to Early Jurassic. In the Causses, these latest are prevalent but in Quercy (France), Poland, Italy, USA, they are also associated with Omithopoda, Thyreophora and Sauropoda ichnites. Footprint areas considered here were generaly under an arid climate. Animals that passed by were heavy and bulky possible Megalosaur trackmakers, and lighter and slender Coelophysids or Ceratosaurs. For all, these areas were pathways as the orientations of the trackways seem point out. The directions followed by these reptiles were without any important variation during the Hettango-Sinemurian stages. These areas were also used from time to time by Crocodilomorpha and may be tetradactyl (I-IV) bipedal avian Theropods. However, the number of such trackways in sites, sometimes substantial, should not lead us to overestimate the trackmakers populations. These last were probably relatively moderately abondant in this inter-supratidal swamp environment. In the Causses, ichnites are connected with former algo-laminated deposits (Algal mats) which were rapidly hardened by means of calcitisation of cyanobacteria. The result has been a moderate depth of footprints; autopodia disturbing only a few cm of the carbonate substrate. Other fossils have been discovered : invertebrates with thin bivalve and gastropod shells, crustaceans tests and plants. These latter suggest the existence of paleomangroves like environments but also continental vegetation periodically overruning the swamp environment during regression/transgression cycles. At these times, wooded parts of it, could become protecting, feeding, resting and nesting places.

  

Le présent volume est l'aboutissement d'un projet né il y a presque cinq ans. En décembre 1991, l'un d'entre nous (MG) prenait des contacts en vue de proposer un symposium sur les mammifères fossiles, dédié à D.E. Russell, dans le programme du 4e Congrès de la European Society for Evolutionary Biology. Ce congrès, baptisé "Evolution 93", devait se tenir à Montpellier en août 1993. Son Comité d'Organisation, animé par F. Catzeflis, recherchait des organisateurs de symposiums. L'idée fut acceptée avec enthousiasme par le second d'entre nous (PDG), et le titre de notre Symposium fut précisé: " Palaeobiology and Evolution of Early Cenozoic Mammals - A Symposium in Honor of D.E. Russell". Le projet fut formellement accepté par le Comité d'Organisation en avril 1992. 

The earliest cetaceans are found in the early Eocene of Indo-Pakistan. By the late middle to late Eocene, the group colonized most oceans of the planet. This late Eocene worldwide distribution clearly indicates that their dispersal took place during the middle Eocene (Lutetian). We report here the first discovery of a protocetid fossil from middle Eocene deposits of Senegal (West Africa). The Lutetian cetacean specimen from Senegal is a partial left innominate. Its overall form and proportions, particularly the well-formed lunate surface with a deep and narrow acetabular notch, and the complete absence of pachyostosis and osteosclerosis, mark it as a probable middle Eocene protocetid cetacean. Its size corresponds to the newly described Togocetus traversei from the Lutetian deposits of Togo. However, no innominate is known for the Togolese protocetid, which precludes any direct comparison between the two West African sites. The Senegalese innominate documents a new early occurrence of this marine group in West Africa and supports an early dispersal of these aquatic mammals by the middle Eocene.
  

A new European rodent from the middle Eocene of Spain, Zamoramys extraneus n. gen., n. sp., appears to be closely related to the middle Eocene chapattimyid rodents of Indo-Pakistan. This contradicts the generally accepted paleobiogeographic hypothesis of a Tethyian barrier between Europe and Asia isolating Europe during the middle Eocene. Because of this barrier, some authors have proposed that European and Asian rodents were not closely related, their similarity being the result of morphological convergence. Here monophyly has been tested, using the parsimony criterion, based on an analysis of dental characters (including discussing of homology and the validity of some characteristics). Our results indicate a phylogenetic relationship among the Asiatic Ctenodactyloidea, Zamoramys from Spain, and the European endemic Theridomyoidea. We also conclude from our analysis that theridomyoids and European ischyromyoids are probably not closely related phylogenetically. 

The Mont-Hélène's fauna [Pyrénées-Orientales, France], includes 15 species of rodents with a new one, Occitanomys montheleni n. sp. among the 9 species of the Murids which are listed. The uncommon Cricetid, Blancomys neglectus, is well represented in the fauna. Peculiarities of the population referred to Slephanomys cf. donnezaniare discussed. The locality a fissure filling may be the oldest one of Tabianian age known in Southern France. The diversity of the Murids gives evidence of a subtropical climate and of a diversified environment which may be linked to the spreading of the coastal plain following the filling up of the Roussillon Neogene Basin. 

The first good skeletal remains of Homogalax protapirinus from the Wasatchian of the Bighorn Basin, Wyoming, indicate that this primitive tapiromorph was more plesiomorphic in many features than primitive equoids including Hyracotherium. Compared to Hyracotherium, Homogalax more closely resembles Phenacodonta (the closest outgroup of Perissodactyla for which postcrania are known) in various details of articular surfaces, muscle attachments, and proportions of the humerus, manus, and pes.Among known taxa, Homogalax most nearly approximates the plesiomorphic postcranial skeletal anatomy of Perissodactyla. 

The mammalian fauna (represented only by isolated teeth) from the early Cretaceous of Morocco includes a number of non-'therian' specimens, characterized by the antero-posterior alignement of the main molar cusps. This assemblage is very heterogenous, including forms (among them Gobiconodon palaios sp. nov.) that can be related to Laurasian triconodonts and several forms so far known only in this part of Gondwana (among them Kryptotherium polysphenos gen. et sp. nov.). The latter, in particular, exemplifies the diversity of the mammalian dental morphology in the Mesozoic, often largely underestimated. In the last part of this paper, the 'therian' affinity of the 'amphilestids' is discussed, on the basis of the arrangement of molar cusps, the interlocking mechanism and the occlusal pattem. 

Aside from Madtsoiidae, anilioids, and Boidae that were studied previously, the middle Palaeocene of ltaborai (BraziI) has produced Ungaliophiinae ("tropidophiids"), booid-grade snakes incertae sedis, and a possible Russellophiidae (Caenophidia) that are described in the present article. This article is the third and final report on the snakes from the locality. The Ungaliophiinae (Paraungaliophis pricei gen. et sp. nov.) are rare whereas the booid-grade snakes incertae sedis (ltaboraiophis depressus gen. et sp. nov., Paulacoutophis perplexus gen. et sp. nov.) are more frequent. A single vertebra is referred to the Russellophiidae (Caenophidia) with reservation. An update of the whole fauna of snakes from ltaborai is provided. Hechtophis austrinus that was tentatively referred to the erycine Boidae is now regarded as a Boidae incertae sedis. Most snakes from Itaborai are known only from the locality. Astonishingly, only the ailioids Coniophis cf. C. precedens gives possible evidence of interchanges between South and North America. The fauna of snakes from Itaborai, as well as the other Palaeocene faunas of snakes from South America are distinct from those of the Cretaceous and the Eocene of South America; they appear to be more different from the Cretaceous faunas than from those of the Eocene. The fauna from Itaborai is the richest and most diverse assemblage of snakes from the Palaeocene worldwide; it shares only a few taxa with other Palaeocene localities.