Abstract
 The validity of Propachynolophus Lemoine, 1891, supposedly an intermediate between Hyracotherium Owen, 1841 and Pachynolophus Pomel, 1847, has been questioned for a long time. A detailed analysis of features on which this genus is based further supported by a formal cladistic analysis demonstrates that Propachynolophus is not a valid taxon. The type species, “Propachynolophus gaudryi Lemoine, 1891” shall be assigned to Propalaeotherium Gervais, 1849, under the new combination Propalaeotherium gaudryi (Lemoine, 1891). “Pachynolophus maldani Lemoine, 1878”, later assigned to Propachynolophus, typifies the new genus Orolophus, under the binomen Orolophus maldani (Lemoine, 1878). The other referred species, “Propachynolophus levei Hooker, 1994” and “P. remyi Checa-Soler, 1997” are poorly documented, and both species shall be provisionally referred to as “Hyracotherium” levei (Hooker, 1994) and “Hyracotherium” remyi (Checa-Soler, 1997), pending new discoveries.
 
  

A new Propalaeotherium species, clearly distinct from the genus Eurohippus, is described. It is characterized by having a similar size as P. voigti from the German Geiseltal localities (MP 11 to MP 13 reference-level), but differs in several features suggesting a slighty more derived morphology. It presents indeed less brachyodont crowns with less prominent and less elevated cingula, slightly larger relative surface of premolars, and a more marked metaconid splitting on cheek teeth. This new species is unknown from other European localities except the nearby Saint-Martin de Londres locality which has been considered older than the MP 13 level. 

We present here a new updated counting of the perissodactyls of Robiac, the type locality of the MP 16 level of the biochronological scale of paleogene mammals and that of the Robiacian stage of Eocene Land Mammals Ages in Western Europe.
The outcrop of Robiac consists actually of two 500m apart loci, Robiac-Nord and Robiac-Sud, considered of the same age according to the current discriminating power, and is dated from -38,7 MA after the last faunal, magnetostratigraphic and climatic calibrations.
It has yielded a very abundant and rich of 21 taxa perissodactyl fauna, topped by the giant Lophiodon lautricense, last representative of the family Lophiodontidae, of which it is the last proved deposit. The Palaeotheriidae are much diversified with 5 genera and 9 species of "Pachynolophinae", 3 genera and 10 species of Palaeotheriinae. Nine taxa have been defined from Robiac: Chasmotherium depereti n. sp., Palaeotherium castrense robiacense Franzen, 1968, the genus Leptolophus Remy, 1965 with the species L. stehlini Remy, 1965 and L. magnus Remy, 1998, Anchilophus (Paranchilophus) jeanteti Remy, 2012, Metanchilophus chaubeti Remy, 2012, Lophiotherium robiacense Depéret, 1917 and Pachynolophus gaytei n. sp.
The faunal Robiac cenogram with the associated flora testify to a hot, wet and forestal environment, likely corresponding to a short warming climatic phase; the broken up fossil bones should have been carried away and then gathered in swamp areas along the banks of a meandering river.
The swarm of mammals of Robiac, the richest of contemporaneous deposits, has been followed by a drastic drop in perissodactyl diversity at the MP 17A level; a crisis which could have originated in a renewal of the global Eocene cooling. Fons 4, the type-locality of this level, is largely scarcer in perissodactyls and its cenogram testifies to a less diversified fauna, with on the whole smaller species, that likely means a cooler and drier climatic environment; a new perissodactyl diversification occurred but later.
  

The knowledge of the Anchilophini has been lately renewed by the discovery of a rather large amount of new material still largely unpublished. This new material offers the opportunity of a systematic revision of this tribe gathering those of European Eocene Equoidea which bear no mesostyle on upper check teeth and display a heavy trend to the molarization of premolars.

A cladistic analysis has made out two genera, Anchilophus (Paranchilophus included as a subgenus), characterized by a marked lophodonty and the transverse narrowness of the cheek teeth, a rather high hypsodonty, the frequent occurrence of "crochets" and "anticrochets" on the superior ones, and a rather weak molarization of the premolars, opposite to Metanchilophus n. gen. whose cheek teeth are more transversally elongated, less high, less lophodont, with cusps better distinct, enamel thicker and premolars more molarized on the whole.

Three species of Anchilophus are recognized, A. desmaresti, type species of the genus, A. (Paranchilophus) remyi and A. (Paranchilophus) jeanteli n. sp.

The genus Metanchilophus is more diversified with the species dumasi, radegondensis, gaudini (whose a new sub-species M. g. fontensis is defined), depereli, castrensis n. sp. and chaubeti n. sp.

The skull anatomy has been moreover described with several taxa; it brings to light (for all that one can generalize) that Anchilophini were light animals with a slender and elongated snout, a thin zygomatic arch, a rather developed encephalon with an advanced gyrencephaly.

The structure of the nasal opening together with the occurrence of epitympanic sinuses and the molarizing process of the premolars corroborate the attribution of this tribe to the family PalaeotheIiidae. 

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. 

A rear skull from the Quercy Phosphorites is described. It documents a new perissodactyl genus, likely assignable to the family Palaeotheriidae and probably paleogene of age. Owing to the lack of any tooth, this family assignment remains however somewhat hypothetical. The specimen is firstly characterised by the presence of a wide epitympanic sinus swelling and hollowing the squamosal shell and the post-glenoid process. This cavity might make up a peculiar pattern of improvement for the hearing apparatus by carrying out a kind of drum near the middle ear, whereas the bony tympanic remains barely bulged and forms but a few developed auditory bulla. This pattern appears an outcome of a trend observed with many palaeotheriids, such as Plagiolophus. Furthermore, the endocranial cast shows a rather high degree of gyrencephaly for a paleogene mammal. The prominent lambdoidal crest points out a powerful nape musculature and a lowered head bearing. Consequently, it is assumed that such an animal was probably living in somewhat open places, at the expense of rather tough vegetables. It might have been accordingly provided with a semi-hypsodont, cement covered, "plagiolophoid" dentition. 

A strong lophodonty, an extreme heterodonty, some hypsodonty and regular overlayings of coronal cement are prominent features of the genus Leptolophus (Palaeotheriinae = Palaeotheriidae s.s.). The histological pattern of the teeth unusually joins type II enamel prisms, characteristic of advanced ungulates, together with archaic features, such as an almost complete lack of Hunter-Schreger zonation and a weak expanse of peritubular dentine. The skull is narrow and slender, with an elongated ante-orbital facial region, a moderately notched nasal aperture, a rather elongated post-canine diastem, parallel zygornatic arches and a fairly dorsally located squamoso-mandibular joint.The functional analysis brings to light "ectolophodont" masticatory cycles with two phases, in which maximum power was applied, contrary to equíds, on hindmost teeth; likewise, skull accomodations to increasing height of the teeth are quite different. This study leads to the assumption that Leptolophus may have been light mammals, living in rather open surroundings, browsing on herbaceous plants or leaves cropped close to the ground. Moreover, it appears that it could have been some inadequacy of dental structures to the dietary, which leaded to quick wear of the teeth and to many enamel notches, but had been somewhat balanced by the early increase of hypsodonty, not induced in such a case by a biotop deterioration (as it will happen at the end of the Eocene). This ínadaptation might account for the short duration of the genus Leptolophus, whose the 3 species, L. stehlini, L. nouletí and L. magnus n. sp. are indeed confined in the level MP 16. Its geographical spreading (as far as known, South of western Europe) and the morphological pattern of its dentition suggest that this genus would have been related to early upper Eocene endemic spanish forms.





  

The St-Dézéry local fauna (3 reptile-, 4 mammal species) is approximately of the same age as the La Débruge or the Ste-Néboule faunas. It conduces to a better dating of the limestones underlying the Célas sandstones. A large part of a mandible of Amphimeryx was found there, which documents the record of this family of small artiodactyls 

The skull remains referred to the genus Palaeotherium are recorded and described. Biometrical tests are made to elucidate intrageneric allometric relationships and to allow comparisons with various other perissodactyls. Apart from the well known shortness of post canine diastems and deepness of the narial opening, the genus is characterized by a great lengthening of the splanchnocranium, owing to a spreading of the post-orbital facial region, by a reduced area of the eye-socket and by the prevalence of the temporal muscle with regard to the masseter; this original shape of the masticatory apparatus needs to be related to the morphology of the jugal teeth and particularly to their  asymmetrical semi-hypsodonty.
    These animals, whose running ability was evidently poor, appear to have been adapted to rather closed environments, feeding on relatively soft vegetable matter; olfactory sense was likely to play a leading part in interindividual and environmental relationships. Such evolutionary trends might explain the disappearance of most of them, as clirnatic conditions deteriorated at the end of the Eocene, before the "Grande Coupure" which affected mammalian faunas at that time.
      Although the present paper is not directly concemed with phylogenetics, it invalidates the supposed ancestor-descendant relationship between P. castrense and P. magnum, and it suggests a possible emergence of the P. medium lineage from a P. siderolithicum stock. Moreover, the structure of the post-orbital facial area allows the  erection of a new sub-genus, Franzenitherium, for the  species lautricense and duvali.

  

The Laprade fauna is chronologically situated between those from Egerkingen and Lissieu and consequently, is close to the MP 14 reference-level of the European mammalian biochronological scale (Symposium of Mainz, 1987).
This new fauna is presently the oldest known in the Quercy phosphorites, formerly the Le Bretou fauna (MP 16) was considered as the oldest one. The Laprade fauna includes 21 species which belong in 7 mammalian orders (Marsupialia: Amphiperatherium bastbergense, Amphiperatherium goethei; Apatotheria: Heterohyus (Gervaisyus) pygmaeus nov. subgen., nov. sp.; Lipotyphla: Saturninia cf. mamertensis, Saturninia cf. intermedia; Chiroptera: Vespertiliavus lapradei nov. sp.; Rodentia: Protadelomys cf. lugdunensis, Elfomys nov. sp.; Primates: Nannopithex cf. filholi, cf. Pseudoloris or Pivetonia; Perissodactyla: ?Palaeotherium ?castrense, small-sized Palaeotherium sp., Plagiolophus sp., Anchilophus sp.; Artiodactyla: Dichobune cf. robertiana, Mouillacitherium cartieri, Tapirulus cf. depereti, Mixtotherium priscum, Pseudamphimeryx schlosseri, and Artiodactyla indet). Sixteen of these species are mentioned for the first time from the Quercy faunas.
The recognition of a new apatemyid, Heterohyus (Gervaisyus) pygmaeus nov. subgen., nov. sp., attests to an early origin of a lineage known in the Late Eocene. The emballonurid bat Vespertiliavus lapradei nov. sp. is presently the earliest record of this genus and family. This Auversian fauna leads to discuss the age of taxa showing archaic features. These taxa were defined on specimens collected in the Quercy during the last century and have never been found in the Quercy localities recently investigated. This fauna bears also evidence of a karstic filling episode older than those previously dated by fossils in the Quercy Jurassic. 

Biographie et liste des publications de S. Garimond. 

La poche à phosphorite de Sainte-Néboule (Lot) a livré au cours des récentes campagnes de fouilles effectuées dans le cadre de la Rep 311 une douzaine de dents ou fragments de dents de paléothéridés à rapporter à 3 taxons. 

Le Quercy est aujourd 'hui un vaste plateau calcaire, parcouru par un réseau karstique actif, profondément
entaillé par des vallées aux falaises abruptes, comme celles du Lot ou du Célé. Sur un sol peu épais domine la forêt de chênes, accompagnés de cornouillers, érables, genévriers. La faune est pauvre, peu diverse, et les nombreux chasseurs se satisfont de gibier d'élevage ...

Il y a trente cinq millions d'années environ, le paysage était bien différent. La période de l'Eocène supérieur, qui s'achevait, avait été chaude et humide, si l'on se réfère à la fois aux paléotempératures (calculées à partir de sédiments marins extra·européens) et aux restes fossilisés de végétaux typiquement tropicaux. 

Deux types de remaniements post mortem me paraissent caractéristiques de l'état de conservation des dents de mammifères fossiles dans les Phosphorites du Quercy :

1) Des destructions localisées d'origine biologique, sous forme de galeries de morphologie très variable creusées dans la dentine et le cément, et impliquant sans doute la participation de différents types de micro-organismes. Ces altérations se sont développées peu de temps après la mort, avant la fossilísation proprement dite et se sont rapidement arrêtées après l'enfouissement dans le sédiment phosphaté.

2) Des perturbations dans les structures de la dentine liées aux variations locales de minéralisation, provoquées par une imprégnation diffuse des zones les moins calcifiées par divers minéraux et probablement surtout de l`apatite. 

The genus Pachynolophus, one of the poorest known of the Palaeotheriidae, includes the brachyodont forms with reduced and non-molariform premolars and with upper molars lacking a mesostyle. Quantitative characters (divers surface indications and elongation of the teeth), while demonstrating a close relationship to Hyracotherium, permit a better differentiation of the genus, confirm its specific splitting, and permit the distinction of three lineages. The skull from Memerlein is taken as the type of a new species, P. Iavocati, of which the dentition is extremely characterized by its lophiodonty, the strong reduction of the premolars and the reduction of the cingula. This characterization testifies to a late age which extends the existence of the genus quite near to the Eocene-Oligocene limit. Compared with the only two skulls known of related species (Hyracotheríum vulpiceps and Pachynolophus Iivinierensis), that from Memerlein is distinguished by progressive characters affecting diferent regions but most particularly the braincase; it is not possible, however, to isolate within this evolution the part which leads to a systematic differentiation. Modernization is translated by a considerable increase in size of the braincase, principally in the frontal region, a development of the facial region with anterior displacement of the dental series and a greater specialization of the masticatory apparatus. This evolution parallels the history of the Equidae of the North American early Tertiary, but certain particularities, the form of the alisphenoid, the presence of an anterior frontal foramen, and the structure of the paroccipital apophysis, testifies to the independance of the European forms. 

The locality of Fons 1, one of the fossiliferous outcrops in the late Eocene limestones of Fons-outre-Gardon (Gard), has yielded varied remains of mammals. The specimens were prepared by dilute acetic acid attack on the rock and by impregnation with an acrylic resin.
This fauna, rich with about 20 species, can be included in the biochronologic zone of Euzet. The Palaeotheridae (sensu Simpson 1945), represented by 6 species, occupy a predominant place; Plagiolophus annectens is particularly abundant, comprising 55% of all the specimens found.
The abundance of this species has permitted a detailed comparative study with the corresponding form from Euzet. The quantitative tests brought out a slight but highly sígnificant difference between the average dimensions of the teeth in the two populations. Moreover and in spite of the feeble rate of evolution in the Plagialophus dentition, diverse archaic particularities can be seen which permit the conclusion that the form from Fons 1 is a little the older. This conclusion is corroborated by observations on the other palaeotherids, particularly Lophiotherium aff. cervulum, whose dental evolutionary stage is intermediate between those of the corresponding forms known from Robiac and Euzet.
A new species of Pachynolophus is described, P. garimondi n.sp., the last representative known of the genus. lts teeth are distinguished by their size, a great heterodonty, and the weakness
of their cingula.