The hipparions from the Dytiko localities of the lower Axios valley (Macedonia, Greece) are studied. The material comes from three localities Dytiko-l, 2, 3 (DTK, DIT, DKO), which are situated near the village of Dytiko, about 60 km northwest to Thessaloniki. Three species have been determined, the medium-sized H. mediterraneum, the small-sized H. matthewi and the very small-sized H. periafricanum. The determined Hipparion species, their morphological characters and their comparison with the other Axios valley material indicate a Late Turolian age for the Dytiko localities. 

The stapes is the last of the middle ear ossicle chain and the smallest bone of the mammalian skeleton. Because it rests on the membrane of the fenestra vestibuli during life, the stapes may often fall within the bony labyrinth cavity when soft structures decay after death. In this work, we highlight the unexpected role that the bony labyrinth plays in the preservation of the stapes. Systematic investigation of the bony labyrinth of 50 petrosal bones of extinct and extant artiodactyls led to the discovery of eight cases of “intralabyrinthine” stapes. Three dimensional reconstructions of these stapes allow documenting stapes morphology of four extinct artiodactyl taxa: Microstonyx erymanthius (Suidae), Elomeryx borbonicus (Hippopotamoidea), ?Helohyus plicodon (Helohyidae), and an undetermined Cainotheriidae; and four extant ones Choeropsis and Hippopotamus (Hippopotamidae), and Tayassu and Phacochoerus (Suoidea). ?Helohyus plicodon from the Middle Eocene documents the oldest stapes known for the order Artiodactyla. Morphological study and metric analyses of our sample of artiodactylan stapes show that stapes are likely to carry relevant phylogenetic characters/signal within artiodactyls, and a potential Euungulata signature. 

The present study of Artiodactyla from the Middle Eocene of the Geiseltal lignite beds concems systematics, biostratigraphy, and palaeoecology on the basis of 174 specimens: isolated remains to more complete skeletons. Instead of the formerly known five species of two families are now recognized 14 species of the Diacodexeidae, Dichobunidae, Cebochoeridae, and Haplobunodontidae. New species are Aumelasia maniai, Anthracobunodon neumarkensis, Masillabune franzeni. Four species of the Geiseltalfauna are definitely known from elswere, and five species are closely related to those from other European localities. Evidently the faunal situation of artiodactyls during the Middle Eocene of Europe was largely uniform. The distribution of artiodactyls within the sequence of the Geiseltal strata corroborates the biostratigraphical concept of the land mammal age Geiseltalian (Franzen & Haubold l986b) as well as the mammalian reference levels MP 11-13 (Franzen 1987). Reconstructions of the skulls and skeletons allow conclusions on the functional morphology and palaeoecology of the artiodactyls of the European Middle Eocene 

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.

  

Micro-CT imaging is an increasingly popular method in paleontology giving access to internal structures with a high resolution and without destroying precious specimens. However, its potential for the study of hypoplasia defects has only recently been investigated. Here, we propose a preliminary study to test whether hypoplastic defects can be detected with micro-CT (μCT) scan and we assess the costs and benefits of using this method instead of naked eye. To do so, we studied 13 fossil rhinocerotid teeth bearing hypoplasia from Béon 1 (late early Miocene, Southwestern France) as positive control and 11 teeth of the amynodontid Cadurcotherium (Oligocene, Phosphorites du Quercy, Southwestern France), for which enamel was partly or totally obscured by cement. We showed that all macroscopically-spotted defects were retrieved on 3D reconstructions and selected virtual slices. We also detected additional defects using μCT scan compared to naked eye identification. The number of defects detected using μCT was greater in the Cadurcotherium dataset (paired-sample Wilcoxon test, p-value = 0.02724) but not for our control sample (paired-sample Wilcoxon test, p-value = 0.1171). Moreover, it allowed for measuring width and depth of the defects on virtual slices (sometimes linked to stress duration and severity, respectively), which we could not do macroscopically. As μCT imaging is both expensive and time consuming while not drastically improving the results, we recommend a moderate and thoughtful use of this method for hypoplasia investigations, restricted for instance to teeth for which enamel surface is obscured (presence of cement, uncomplete preparation, or unerupted germs). 

The present study describes in detail the anterior limb osteology of a molossid chiropteran of the genus Tadarida, from Céreste, a Stampian locality in the Apt-Forcalquier Oligocene basin already known for its fishes, plants and insects.

A comparision with the Miocene forms T. srehlini from Saint-Géraud localities and T. sp. from Württemberg, also with the recent forms T. teniotis and Eumops perotis, does not show any clear morphological differences between the Tertiary and Recent Tadarida, indicating a rather noticeable anatomical stability, not exceptionnal indeed among Chiropterans. The Céreste fossil exhibits however slightly primitive wing proportions if compared to the Saint Gérand Aquitanian species.

Several remarks deal with the peculiar relationships between the ecology of the molossids and their kind of fossilisation, frequently associated with sedimentary facies of the lacustrine type.
  

Morphological and biometrical description of skulls, teeth, and limb bones of extant and fossil Old World herniones (including E. hydruntinus) and of New World 'stilt-Iegged' and other slender species from Blancan to Holocene. An Appendix presents ways in which the approximate size of some missing bones or dimensions may be deduced from available ones.

The discussed and/or illustrated fossils were found in Bolivia (Tarija), Canada (Yukon), China (Choukoutien, Gulongshan, Jiling, Loufangzi), Ecuador (Oil Fields), Ethiopia (Melka Kunturé), France (Lunel-Viel), Germany (Süssenborn), Greece (Agios Georgios, Petralona), Hungary (Dorog), Italy (Romanelli), Mexico (Cedazo, San Josecito), Mongolia (Sjara-osso-gol), Spain (Venta Micena), ex-Soviet Union (Akhalkalaki, Binagady, Chokurcha, Chukochya, Kabazi, Kolyma, Krestovka, Kurtak, Staroselie, Tologoj), USA (Alaska, Arkalon, Cedar Meadow, Channing, Conkling, Dry Mountains, Hay Springs, Leisey Shell Pit A, Lissie Formation, Natural Trap, Pool Branch, Powers Ranch, Rock Creek, San Diego, Santo Domingo, Seymour Formation, Shelter, Slaton, Trinity River). Numerous raw or statistically elaborated data are given in Tables.

There is no evidence for the existence of Old World hemiones in the New World nor of 'stilt-Iegged' equids in the Old World. The first 'stilt-Iegged' equid was found at Santo Domingo, New Mexico, and is believed to be Late Blancan. It was probably at the origin of E. calobatus (Arkalon, Rock Creek) and of the smaller E. semiplicatus (Channing, Rock Creek). Slender, but not 'stilt-Iegged', equids found at Natural Trap, Wyoming, ca. 12 ky ago belong to Amerhippus. AlI these species share with Oid World Sussemiones (and some hemiones) peculiar patterns on the lower cheek teeth.

The slender Equus sp. B of Leisey Pit A, Florida, ca. 1.2 Ma, as weIl as Amerhippus francisci and E. tau (probably a senior synonym of E. quinni) share conventional lower cheek teeth patterns. The skulls of A. francisci and E. tau, however, are quite different.

Paleontological data suggest a common origin of Amerhippus, Sussemiones, and 'stilt-Iegged' equids during the late Blancan. Old World hemiones seem to have differentiated later. 

Abstract: A very tiny cuckoo, Chambicuculus pusillus, was recently described after a few fragments of tarsometatarsi from the late Early/early Middle Eocene of Djebel Chambi, Tunisia. New remains, notably a coracoid, confirm the attribution of this genus to the recent family Cuculidae. This coracoid shows a very elongate and strap-like processus procoracoideus. This morphological feature is otherwise only known in the family Cuculidae. The characteristics of the coracoid and tarsometatarsi show that Chambicuculus is morphologically more advanced over the other stem cuculids described in Europe and North America. Chambicuculus is the oldest Cuculidae known so far. 

Based on fossil discoveries and phylogenetic studies, an Eocene Asian origin for hystricognathous rodents and anthropoid primates has gained strong support in recent years. The two groups then invaded both Africa and South America, which promoted their evolutionary success. However, the fossil record has so far failed to constrain the nature and precise timing of these pivotal dispersal events. In Africa, given the apparent absence of hystricognaths and anthropoids in early to early middle Eocene localities, it is suggested that these mammal groups dispersed from Asia to Africa sometime during the middle Eocene. In this paper, we report the discovery of several isolated teeth of a rodent from a new vertebrate locality situated in central Tunisia (Djebel el Kébar, KEB-1), dating from the late middle Eocene (Bartonian, ~39.5 Myr). These fossils document a diminutive new species of Protophiomys (P. tunisiensis nov. sp.), a basal genus of hystricognathous rodents which is well known from several North African mammalian-bearing localities of the end of the Eocene. The teeth of P. tunisiensis display a suite of anatomical details comparable with those observed in the other species of the genus, but with a lesser degree of development. Such an apparent primitive evolutionary stage is corroborated by the greater antiquity of this Tunisian species. P. tunisiensis nov. sp. is so far the most ancient representative of hystricognaths in Africa. However, it can be expected that hystricognaths were already present on that landmass given the new data on early caviomorphs recently reported from South America (at ~41 Myr). The arrival of hystricognaths in Africa from South Asia certainly predates the depositional period of the Kébar sediments, but perhaps not by much time. 

Redécouvertes en 1968 dans le bois du Bretou, les deux poches à phosphorites qui ont livré cette faune sont parmi les plus méridionales du plateau quercynois. De faible profondeur, huit mètres, les cavités sont de petite taille et font partie d'un complexe de sept fissures réparties dans les bois du Bretou. Les autres trous n'ont livré aucun fossile, soit parce qu'ils avaient été intégralement vidés de leur sédiment, soit que des venues d'eau en interdisent l'accès.
Dans les deux poches fossilifères, proches de quelques mètres, le sédiment que l'on rencontre en placage sur les parois présente un aspect particulier. Il s'agit en effet d'une brèche très indurée, où la calcite domine, et contenant assez peu de sédiments sidérolithiques. Les ossements contenus dans cette brèche sont assez nombreux, très souvent brisés. Les dents isolées y sont relativement rares. Exploitées séparément, les deux poches ont livré des faunes identiques, aucune différence chronologique ne pouvant être décelée entre elles. Les formes rencontrées dans une seule des cavités sont rares mais la poche la plus riche est celle dénommée Le Bretou 2.
L'intérêt de la faune récoltée réside dans son ancienneté. On sait en effet que H.G. Stehlin avait signalé dans les collections du Quercy des restes attribuables à Lophiodon lautricense, mais la provenance exacte de ces fossiles reste inconnue. Au Bretou si nous n'avons pas rencontré à ce jour le Lophiodon, il n'en demeure pas moins que l'âge de la faune récoltée doit être proche de celui de la faune de Robiac. 

Of the ten cricetid species from the Oligocene of Western Europe, attributed until now to the genus Eucricetodon, only four prove to be utilizable - E. atavus, E. huberi, E. praecursor, E. collatum - to which it is possible to add two forms newly described: E. huerzeleri and E. quercyi. The evolullon of the genus Pseudocricetodon is also the subject of new observations. The study of the dental morphology allows us to distinguish in these two genera three lineages beginning in the middle Oligocene:

- Lineage P. montalbenensis-P. thaleri (from Montalban to Goderet), of small size, without
increase in size.
- LineageE. atavus-E. infralactorensis (from Hoogbulsel to Estrepouy), of middle size, with a regular increase in size.
- Lineage E. huerzeleri-E. haslachense (level of Montalban to that of Estrepouy), of large size, with an increase in size.

Two other forms are equally represented in these loealitles: Heterocricetodon aff. helbengi and Melissiodon quercyi. It has been possible to attribute a precise age (zone of "Cournon") to the last species, which has been defined by Schaub (l925) from material in the Old Quercy collections. The genera Pseudocricetodon, Eucricetodon, ? "Cricetodon", Leydimys, Eumys, differentiated at the beginning of the middle Oligocene in Europe, Asia and North America, seem to derive from a common ancestral group. The place of origin of this group could be situated in Asia. 

A new hypothesis for the phylogenetic relationships of recent anomalurids and graphiurids is proposed, based on information from evolutionary lineages of Paleogene European rodents, particularly Gliridae, and Eocene Algerian Zegdoumyidae. Differences in first occurrences, in paleogeography, and in infraorbital structure in glirids (protrogomorphy and pseudomyomorphy) and graphiurids (hystricomorphy) separate Graphiuridae from Gliridae (Graphiurinae is here raised to family rank). Similar considerations, and dental morphology, suggest that Anomaluridae (appearing in the late Eocene) and Graphiuridae (appearing in the Pliocene) are related to early Eocene Zegdoumyidae. 

A large avian femur recently discovered at the Late Cretaceous Montplo-Nord locality at Cruzy (Hérault, southern France) is referred to the giant bird Gargantuavis philoinos. The estimated mass of the bird is 57 kg, within the range of living cassowaries. The specimen provides new evidence about the anatomy of G. philoinos, notably showing that the distal end of the femur was similar to that of modern birds in having a condylus lateralis subdivided into two semicondyles. A new diagnosis of Gargantuavis philoinos is provided and the taxon is placed in a new family of basal ornithurines.
  

A new carcharhinid shark species, Physogaleus hemmooriensis sp. nov., is described from the Lower Hemmoorian (Behrendorfian, late Burdigalian, early Miocene) of Werder, Lower Saxony, Germany. P. hemmooriensis also occurs in the Edegem and Antwerpen Sands Members of the Berchem Formation, Belgium, and in the Miste Bed, Aalten Member of the Breda Formation, The Netherlands, which have an early to middle Miocene age. In the Western Atlantic region, the taxon is present in the early Miocene Calvert Formation of Delaware, U.S.A, which is largely contemporaneous with the Hemmoorian. 

Prolagus sardus is the last representative of the diverse lineages of European endemic ochotonids. It is also the most abundant in the collections. The previous studies made of this species have established rather well its dental morphology, its phylogenetic position, its geographic and temporal distribution, and its intraspecific individual variation. On the other hand, no osteologic study has fully utilized the superb material from Corsica and Sardinia  collected by Forsyth Major.
Nearly all of the parts of the skeleton are represented in this material; they are here described and figured. Comparisons are made with Ochotona, the only surviving genus of the family, as well as with the living leporids, in particular Romerolagus, Oryctolagus and Lepus. A brief examination of the dentition reveals some particular characters of the incisors. The osteologic study allows limited interpretations to be advanced concerning the posture, mastication. locomotion and some other adaptive features of P. sardus.
P. sardus appears as an incontestable ochotonid, but it differs from Ochotona by some characters which are found in leporids. Certain of the characters are here judged primitive for lagomorphs. The genus Ochotona therefore can no longer be considered as the image of a primitive lagomorph, because in spite of the lack of cursorial adaptations which differentiate it from leporids, it presents other specializations which are common to it alone. 

Pleistocene bird fossils have been studied from nine localities on Crete. Part of this material was described earlier by the author (Weesie, 1982) and will not be treated here in extenso, the results will be incorporated. More than one third of the over 10,000 fossil bird bones available could be identified ; they were found to represent at least 65 bird species. The following species of the Pleistocene Cretan avifauna are new to the fauna of Crete : Branta ruficollis, Haliaeetus albicilla, Gyps melitensis, Aquila chrysaetos simurgh n. ssp., Ketupa zeylomensis, Aegolius funereus, Dendrocopos leucotos, Zoothera dauma, Turdus iliacus and Pyrrhula pyrrhula. The Pleistocene Cretan avifauna differs less from comparable mainland avifaunas than (fossil) avifaunas from oceanic islands do. Still, the Pleistocene Cretan avifauna has two qualities that are characteristic of island avifaunas : the almost complete absence of a group of birds (the Galliformes) and the presence of two endemic (sub)species : the giant eagle Aquila chrysaetos simurgh n. ssp. and the long-legged owl Athene cretensis (Weesie, 1982). The new subspecies is described in the present study.
These endemic birds of prey were found in association with their supposedly principal prey species (now extinct as well) : endemic mice for the owl and endemic deer for the eagle. Endemic mammals have been found in association with endemic birds of prey on many islands, not only in the Mediterranean. There is evidence that the size of endemic birds of prey becomes optimally adapted to their feeding on certain endemic mammals, especially rodents. Another characteristic of the Pleistocene Cretan avifauna is the great number of species of birds of prey. This appears to be a common characteristic of fossil avifaunas from caves on Mediterranean islands as well as from caves on the European mainland. However, we think that ecological conditions on Pleistocene Crete (especially the abundant presence of mice) helped to account for the high representation of birds of prey. Furthemore, the fossil avifauna enables us to draw some conclusions about the climate and vegetation on Pleistocene Crete : it is concluded that the climate was cooler than today and that Crete was largely covered with forests. Finally, the reasons for the extinction or disappearance from Crete of some bird species of the Pleistocene Cretan avifauna are discussed. 

The analysis of oxygene isotope variations as well as paleobotanical data suggest that the Oligocene/Miocene boundary corresponds to a transitional period marked by floristical and climatic variations. During this period, the pyreneo-alpine tectonics has contribued to modify the geography and western Europe landscapes. Faunal changes (appearances, extinctions, migrations) are observed in different mammalian groups, notably in the rodents. A study of the evolutionary trends and patterns in paleogene rodents is involved for the period ranging from level MP 28 of the Late Oligocene to the Early Miocene, including the Oligo-Miocene boundary.
The Rodents fauna from the sites of Venelles (Bouches-du-Rhône District, France) and Thezels (Lot, France), previously mentionned in litterature, have been studied. The first description of the Eomyidae of La Milloque (MP 29) has been completed. These faunas are compared to those from various localities dating from the considered period. In La Milloque, a new representative of the Eomys species is described next to a form close to Rhodanomys hugueneyae ENGESSER, 1987. It is the Eomys milloquensis nov. sp., the likely descendant of Eomys quercyi COMTE & VIANEY-LIAUD, 1987. Two new species are also described in Thezels: Eucricetodon thezelensis nov. sp., resulting from a likely and local evolution of Eucricetodon praecursor (SCHAUB, 1925) from La Milloque, which, in the same geographic area, could be at the origin of Eucricetodon hesperius ENGESSER, 1985 from Paulhiac. Plesiosminthus admyarion nov. sp., quite distinct from Plesiosminthus schaubi VIRET, 1926, which announces Plesiosminthus myarion SCHAUB 1930. Venelles 'Plesiosminthus schaubi population is considered as a sub-species, named Plesiosminthus schaubi meridionalis nov. subsp. New phylogenetic patterns are proposed. Among the Eomyidae, a quantification of various features of the M1-2/ crown (hypsodonty, degree of abrasion, occlusal angle, state of development of the I and V anticlines), and a comparison with the occlusal diagram of the other teeth among various other populations allows a more efficient separation of Eomys and Rhodanomys genera. In Western Europe, and within this period, it finally does not seem possible to gradually connect the genus Eomys to the genus Rhodanomys. The evolution of the Eomys quercyi - milloquensis lineage seems to underline a similar evolution to that which may have led from the Eomys to the Rhodanomys form. The latter which appears totally accomplished at level MP 29 of the Oligocene is considered as an immigrant. If we compare the most representative species of the Venelles, Thezels, and Coderet sites, (i.e. Rhodanomys, Eucricetodon, Adelomyarion, Peridyromys, Plesíosminthus), it becomes impossible to confirm their biochronological separation. The noticeable differences between the populations may be interpreted as geographical variations. An explanation to these variations, and to fauna's evolution during the Late Oligocene and Early Miocene can be found in the environmental modifications, supported by isotopic, paleobotanical and sedimentologic analysis. A tentative reconstruction of the environments is attempted by the cenogram method. The analysis of the fluctuations of fauna's diversity shows variations which may be correlated to a drop in temperature at MP 29, during the Late Oligocene, followed by an increase in temperature along with an aridity phenomenom, during the basal Miocene (MN O).The confrontation of various methods give the opportunity of reconstituting and comparing the evolution of the environment of three sequences of sites chosen from different regions. Ecological affinities of various rodents' species are being examined. It is possible to consider that the integration of all the conclusions resulting from this study should lead to an explanation to the evolution of rodents for the period around the Oligocene-Miocene boundary. The site of Coderet- level 3- would be posterior to the latter, at the beginnig of the Miocene, and would mark the level MN 0 of the Aquitanian.

  

Anthracotheres from the Pondaung Formation, Myanmar, are considered as one of the most primitive artiodactyl groups and they represent the oldest known record in the world. Thus, the understanding of this group has numerous implications for evolutionary biology and biochronological correlations. However, the systematlcs of these mammals has been interpreted in different ways, and the main debate focuses on the number of taxa represented in the Pondaung Formation. The revised taxonomy proposed here is mainly based on the relative development of the upper molar W-shaped ectoloph, system of crests and stylar cusps, and on body size. On the basis of these characters, they are classified into four genera including six different species. Two well-known genera, Anthracotherium and Anthracokeryx, are validated and more precisely diagnosed. Anthracokeryx possesses a better developed W-shaped ectoloph, system of crests and stylar cusps than Anthracotherium, which displays notable differences with the more derived representatives of this genus. Both of these Pondaung genera show evidence for sexual dimorphism. However, the incompleteness of fossil material fueled a debate concerning the status of two additional Pondaung anthracotheres, Siamotherium and Anthracohyus. The latter genus is of uncertain affinities, but it has been considered as a hippopotamid ancestor. Despite new material attributed to these two forms, additional discoveries are still required to establish their taxonomic status. The hypothesis that Southeast Asia was the centre of origin of Anthracotheriidae is supported by the retention of numerous primitive dental characters in these taxa and by the antiquity of the Pondaung Formation, to which an age of 37 My is now generally accepted. 

There is only one suid known in the Upper Miocene of Montredon (Hérault): Microstonyx (Limnostonyx nov. subgen.) antiquus (KAUP). It is differenciated from Microstonyx major by the presence of upper and lower canines which are considerably longer and biger. Its presence at Montredon corroborates the palustrine habitat for the species. 

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.