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 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.

  

A fragmentary mandible and maxilla of a small sized Agriotherium of a young individual discovered from a Pliocene fissure filling in Xiaoxian county (Anhuei Province, China) are described. Judging from the morphology of the dentition and its dimensions the new material can be identified as Agriotherium inlermedium (STACH l957). Hendey's proposition (1980) that the Agriotherium species are derived from Indarctos is reconsídered on the basis of the new documents. As a result of a more general phylogenetic discussion it can be stated, that: 1. the supposed size increase as well as other trends, leading from Indarctos to Agriotherium are untenable ; 2. there are no positive indications to assume a phylogenetic transition of these two genera. 3. there are no real arguments in favor of an adaptational reversal in the evolution of Agriotherium. Hence, many features of that genus supposed by Hendey to be derived are plesiomorphic ; 4. regardless of the previous points it is methodologícally impossible to establish direct ancestor - descendant relationships between Indarctos and Agriotherium species, as Hendey did. Based on the data available and especially on the characters of the new material from China it is more likely that Agriotherium and Indarctos are two genera which developed independently. While advanced Agriotherium species, e.g. A. africanum, resemble in some respects Indarctos by adaptational analogies, more primitive species, e.g. Agriotherium intermedium, are quite dissimilar to lndarctos. While Indarctos might be derived from an Ursavus like forerunner, Agriotherium has its roots more likely somewhere in between Ursavus and the Hemicyon-group. 

The formic acid preparation of a nodule collected from the Lower Triassic of Spitsbergen, and containing a skull with some associated postcranial elements of Lyrocephaliscus euri (WIMAN), precises and enriches previous descriptions of the reputedly well-known trematosaur. lt permits more detailed descriptions of the occipital region, of the course of the carotid arteries and collateral vessels, and refutes the previous description of a stapedial foramen in trematosaurs. The discovery of pleurocentres associated with the other vertebral components confirms the primitive rhachitomy of trematosaurs.