Abstract: Paleontological exploration in the Turkana Basin near Ileret, Kenya yielded the most complete adult elephant cranium (KNM-ER 63642) known from the late Miocene to mid-Pliocene. KNM-ER 63642 derives from the lower Lonyumun Mb. of the Koobi Fora Fm. and dates to the early Pliocene, >4.3 Ma. The cranium is immense in size and preserves most of its structures including left and right M2-3, permitting its comprehensive comparative study and secure taxonomic assignment to Loxodonta adaurora. Features distinctive of the species and exhibited by KNM-ER 63642 include very elongate, divergent tusk alveoli, a short, biconvex cranial roof, anterosuperior angulation of the occipital planum, non-inflated occipital planum and absence of supralateral parietal "bossing," broad, flat premaxillary nasal processes, broad, laterally downturned nasal aperture superior to the level of the orbits, and M3s with wide, subhypsodont plates that are parallel-faced and separated by U-shaped transverse valleys. The M3s also exhibit characteristic L. adaurora traits of greatest width at their bases, rounded cross-sectional shape, thick enamel, abundant cementum, and strong anterior and posterior accessory conules. Of extant taxa, KNM-ER 63642 most closely resembles crania of African elephants. Its inclusion in the Loxodonta clade is tenuous, however, because shared features are either symplesiomorphic or are difficult to test for synapomorphy due to the poor fossil record of crania of late Miocene-early Pliocene elephants. Overall, the cranial morphology of KNM-ER 63642 is unexpectedly advanced for an elephant of its antiquity. Its anteroposterior compression and height are concordant with efficient proal masticatory action, indicating that by the early Pliocene L. adaurora evolved craniodental adaptations in phase with feeding preference for C4 grasses. The advantage of synchrony of morphology and behavior is reflected by the dominance of the species in the greater Turkana Basin during that interval.
  

The wear facets of primitive rodents can be homologized with those of primitive primates and ungulates. As in primates, the jaw movement was ectental, with an increased anterior component in the lingual phase (phase ll). The buccal phase (phase I) in rodents approaches the horizontal and it tends to be reduced in importance in comparison with the lingual phase. ln more advanced rodents the efficiency of grinding is increased by the development of additional cutting edges of enamel (e.g. enlargement of hypocone, development of mesoloph and lingual sinus). The buccal phase movement becomes lined up with the lingual phase movement to form a single oblique chewing stroke,resulting in planation of the crown. As the stroke becomes more longitudinal (propalinal) the enamel edges become more transverse. In Muridae propalinal chewing evolved before the loss of cusps, facets were reorientated and additional cusps developed. 

New records of terrestrial mammals from the Qasr el Sagha Formation, Fayum Depression, Egypt are reported, and the stratigraphic occurrences of these fossils noted. These include additional specimens of Moeritheríum, Barytherium, and anthracotheres, as well as the oldest record of a hyracoid in the Fayum.These Eocene mammals occur almost exclusively in the alluvial deposits of the Dir Abu Lifa Member of the Qasr el Sagha Formation and show close affinities to the faunas from the lower sequence of the Jebel Qatrani Formation. There is no evidence of a more marked faunal discontinuity between the Qasr el Sagha and Jebel Qatrani Formations than there is across any of the three major breaks in sedimentation that exist within the Jebel Qatrani Formation. The faunal similarities between fossils of the lower sequence of the Jebel Qatrani Formation and of the upper part of the Qasr el Sagha Formation is consistent with recent paleomagnetic dating that suggests that these rocks differ in age by only one to two million years. 

Several endemic deer remains from the Middle Pleistocene deposits of the Castiglione cave (Oletta, Haute-Corse) are examined here. A morphometric analysis allows to relate them to a new insular subspecies Cervus elaphus rossii. The bones were compared with those of the mainland early Middle Pleistocene subspecies Cervus elaphus acoronatus Beninde and the European species Cervus elaphus Linné (Late Middle Pleistocene and Upper Pleistocene forms (continental and insular)). The Castiglione fossil shows peculiar morphofunctional features in its appendicular skeleton suggesting a morphological convergence with certain Bovidae. 

The Paleogene faunal assemblage from Antofagasta de la Sierra (Catamarca, Argentina), is here presented, both in its geological and systematic aspects. The fossil bearing levels are referred to the Geste Formation (Pastos Grandes "Group"). The described specimens belong to the Classes Reptilia (Orders Crocodylia, Serpentes and Chelonii) and Mammalia (three taxa from the Superorder Marsupialia, representatives of the Orders Edentata, Condylarthra, Pyrotheria and Astrapotheria, and six families of the Order Notoungulata). This fauna is referred to the Mustersan Age, which in Patagonia represents the Middle Eocene. Such chronologic assignment is based on the presence of characteristic taxa, their evolutionary stage and on stratigraphic evidence. Finally, a brief comparison with other faunal assemblages from the Early Tertiary of Argentina and Chile, is presented. 

Mammal-bearing localities have been discovered in the marine and lacustrine series of the middle Ilerdian (Lowermost Eocene) from Southem France (Minervois and Corbières). In the localities of Fordones, Monze, Fournès, and La Gasque, thirty mammal species have been identified. Among others, they include ischyromyid rodents (Microparamys and Pseudoparamys), paromomyid and adapid primates (Arcius and Donrussellia), new insectivores, condylarths, and a dyspternine pantolestid. These faunas provide new informations on the early Eocene Mesogean faunas of Rians and Palette. The assemblages of primates and rodents from Fordones support good  correlations with Palette which was recently placed near the standard-level of Dormaal (MP 7). In fact, Palette and Fordones could be even older than Dormaal. Consequently, there seems to be a relatively important temporal gap between the late Paleocene of Cernay and the Sparnacian of Dormaal. This gap could be partly filled with the Mesogean faunas of Palette, Fordones, and Silveirinha. On the basis of these new mammal faunas the marine middle Ilerdian is proved to be older than the Cuisian stage of the Paris Basin. With regards to the position of the Fordones fauna at the top of the NP 10 calcareous nannoplankton biozone, the westem European paleomammalogists Paleocene/Eocene boundary could be situated between the NP 9 and NP 10 biozones. 

Tetraceratops insignis is known from a single, crushed skull from the Lower Permian of Texas. Its unique proportions and osteological details gained central meaning in the question of the origins of Therapsida since this early synapsid has been determined as the oldest and less derived therapsid. Apart from Tetraceratops, the ‘mammal-like’ Therapsida and their sister, the pelycosaur-grade Sphenacodontidae, are separated by one of the longest ghost lineages in tetrapod fossil record. However, the minor, though well justified critique faced insistent publication regarding the therapsid hypothesis. A carefull re-evaluation of the holotypic skull reveals that therapsid traits cannot be supported, including a rejection of the formerly supposed adductor shelf in the temporal fenestra. Increased understanding of ‘pelycosaur’ character variation underlines a haptodontine-grade or, less likely, sphenacodontid position for Tetraceratops. 

An interpretation of the marine reptile fossil record, based on the existing litterature and complemented by the review of ancient collections and the study of new material, permits a better understanding of Mesozoic marine ecosystems. An inventory of the marine reptiles known from the Lower Triassic to the Paleocene is presented: 46 families, about 200 genera and 400 species have been recorded. This data base includes commentaries about systematics, stratigraphical ranges and geographical distribution of taxa. Marine reptiles include a mosaic of not necessarily closely related groups: ichthyosaurs, thalattosaurs, hupehsuchians, pachypleurosaurs, placodonts, nothosaurs, plesiosaurs, pliosaurs but also crocodiles, lizards, snakes, turtles. The diversity studies reveal that the fossil record of marine reptiles has been punctuated by two mass extinctions, during the Middle-Upper Triassic transition and at the Cretaceous-Tertiary boundary. The Jurassic-Cretaceous and Cenomanian-Turonian boundaries (the latter marked by the disappearance of ichthyosaurs) are potential crisis periods, but current data are not sufficient to reach conclusions. The Ladinian-Carnian transition is characterized by the disappearance of 64 % of families and affects essentially coastal forms. This extinction coincides with an important regressive phase. During the Upper Triassic, a faunal reorganisation within marine reptiles leads to the progressive disappearance of near-shore forms and to the development of pelagic groups. During the Maastrichtian-Danian crisis, 36% of families died out. Large-sized pelagic forms such as mosasaurs and elasmosaurs were the most affected and their extinction seems to have been rather sudden. On the other hand, pliosaurs and protostegid turtles became extinct, but were already declining. The survivors were near-shore forms such as crocodiles, snakes and some turtles and they may have taken refuge in freshwater environments. A break in the food chain based on phytoplankton is proposed as an extinction scenario for pelagic forms.

  

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 principles and practices employed in establishment and recognition of South American land mammal ages are reviewed along with previous and present concepts of distinguishing time, rock, and faunal units. Previous chronological arrangements of South American Tertiary land mammal faunas are appraised on the basis of recent geological and paleontological data. Twelve South American Tertiary land mammal ages are here recognized [from oldest to youngest, Riochican (middle to late Paleocene); Casamayoran (early Eocene); Mustersan (middle Eocene); Divisaderan (late Eocene); Deseadan (early [to middle?] Oligocene); Colhuehuapian (late Oligocene); Santacrucian (early Miocene); Friasan (middle Miocene); Chasicoan (late Miocene); Huayquerian (latest Miocene); Montehermosan (early to middle Pliocene); and Chapadmalalan (late Pliocene)]. As all except the Friasian were originally defined on the basis of Argentine faunas, these are discussed first and at length, and each is reviewed with discussion of type locality, stratigraphy, type fauna, and faunal correlations. Non-Argentine faunas are then discussed country by country in alphabetical order.

     A review is given of radioisotope dates obtained on volcanic rocks (i.e., basalts, tuffs) associated with mammalbearing beds in Argentina. Based on these age determinations and on correlation of the late Tertiary land mammals involved in the interchange between North and South America, a chronology of South American land mammal ages correlated with North American land mammal ages and European marine stages is proposed.

It is concluded that South America was an island continent through most of the Tertiary Period (ca 65 to about 3 Ma). As a result, the land mammal fauna of South America developed in isolation and was dominated by autochthonous endemic groups. Toward the end of the Tertiary (i.e., middle Miocene) a unique faunal balance had been achieved by the descendants of the ancient inhabitants (notoungulates, litopterns, condylarths, astrapotheres, edentates, marsupials) and of later (late Eocene) waif immigrants (caviomorph rodents, platyrrhine primates). A prominent feature of this mammal fauna was the combination of carnivorous and omnivorous marsupials with native placental herbivorous ungulates, subungulates, and edentates.

Sometime during the late Miocene, a limited but important interchange of mammalian taxa between North and South America took place. Procyonids (raccoons and their allies), a group of North American origin, first appear in South America in strata of Huayquerian Age, while members of the extinct South American ground sloth families Megalonychidae and Mylodontidae first appear in North America in early Hemphillian time. These groups dispersed along island arcs before the appearance of the Panamanian land bridge in the Pliocene (ca 3.0 Ma). Cricetine rodents, a group of North American origin, are first known in South America in strata of Montehermosan Age. The known taxa are too advanced and diversified to be considered the first of this group to invade South America. lt is believed by some workers that these rodents arrived before the Montehermosan, possibly in the late Miocene or earlier, by waif dispersal from North America.

The isolation of South America ended with the appearance of the Panamanian land bridge, which provided a direct, dry land connection between the two Americas. Across this portal an extensive interchange of terrestrial faunas occurred, and the fossil record documents an intermingling of these long-separated land mammals faunas.

      The beginning of this interchange by land route in South America is marked by the appearance of mammals which evolved from North American emigrants in the Chapadmalal Formation of Argentina. These include a mustelid (Conepatus), a tayassuid (Argyrohyus), and four genera (Akodon, Dankomys, Graomys, Reithrodon) of cricetine rodents. The appearance of this contingent of northern animals favors the existence of the Panamanian land bridge by this time. Likewise, a large number of terrestrial vertebrates of South American origin appear in North America in beds of late Blancan Age date around 2.7 Ma. Among the mammals are Neochoerus, Erethizon, Glyptotherium, Glossotherium, Kraglievichia, and Dasypus. 

Previous chronological arrangements of South American Quaternary land mammal faunas are appraised on the basis of current geological and paleontological data. Three South American late Pliocene-Pleistocene land mammal ages are conventionally recognized, from oldest to youngest, the Uquian, Ensenadan, and Lujanian ; all are defined on Argentine faunas.

     The Uquian is based fundamentally and historically on the fauna from the Uquía Formation in Jujuy Province, northwestern Argentina. Important known formations in Argentina yielding Uquian Age faunas include the sub-surface Puelche Formation (or Puelchense) near the city of Buenos Aires, and the Barranca de Los Lobos and Vorohué Formations between Mar del Plata and Miramar, Buenos Aires Province. A tentative subdivision is propos-ed for the Uquian into three subages based on knowledge of the Mar del Plata-Miramar sequence, from oldest to youngest, the Barrancalobian, Vorohuean, and Sanandresian. In Argentina the Uquian is presently marked by the first known record of Scelidodon, Hydrochoeropsis, Ctenomys, Canidae, Ursidae, Gomphotheriidae, Equidae, Tapiridae, Camelidae, Cervidae, and the last known record of Thylatheridium, Thylophorops, Dankomys, Eumysops, Pithanotomys, Eucoelophorus, Hegetotheriidae, Sparassocynidae, and Microtragulidae.

The Ensenadan Age is based on the fauna from the Ensenada Formation near the city of Ensenada, Buenos Aires Province. In Argentina the Ensenadan is marked by the first known record of Lomaphorus, Neothoracophorus, Plaxhaplous, Cavia, Lyncodon, Lutra, Galera, Smilodon, Dicotyles, Lama, Vicugna, the last known record of Orthomyctera, and the only known record of Brachynasua.

     Typícal beds of late Lujanian Age in Argentina consist of fluvial deposits occupying stream channels, and shallow basins, often incised into beds of early Lujanian (i.e. Bonaerian of early workers) and Ensenadan Age. The Lujanian Age is based on a fauna from beds along the Rio Luján, about 65 km west of the city of Buenos Aires, Buenos Aires Province. The Lujanian in Argentina is marked by the first record of Equus, Chlamyphorus, and Holochilus, and the last record of Megatherioidea, Glyptodontoidea, Arctodus (=Arctotherium), Smilodon, Litopterna, Notoungulata, Proboscidea, Equidae, Morenelaphus, and Palaeolama.

   These land mammal ages are often difficult to recognize in other South American countries. The compositions of South American Pleistocene faunas vary with the environment. Some taxa were widely distributed in fossil deposits throughout the continent, but their occurrences need not reflect synchroneity. This is a result of changing climates and habitats in time. Consequently, proposed intracontinental correlations need confirmation based on magnetostratigraphy and a radioisotope time scale. Paleontologic characterizations of these land mammal ages (i.e. first and last record, and guide fossils) are useful for much of Argentina, but extensions to most of the other parts of South America are at best tenuous.

The majority of known non-Argentine Pleistocene faunas are believed to be Lujanian in age. Possible non Argentine early Pleistocene (Uquian) faunas include Ayo Ayo and Anzaldo in Bolivia, and Cocha Verde in southern Columbia. A possible middle Pleistocene (Ensenadan or early Lujanian) fauna is the Chichense of Ecuador. Paleomagnetic and radioisotopic date (MacFadden et al., 1983) clearly indicate that the greater part of the Tarija fauna (Bolivia) is Ensenadan in age.

  The end of the Pleistocene and beginning of the Holocene in South America is marked by extinction of nearly all large mammalian herbivores and their specialized large predators. Radiocarbon age determinations suggest that large scale extinctions of megafauna occurred between 15,000 and 8,000 yrs. B.P. (years before present). 

A fossil species of the extant genus Leuconoe, L. lavocati n. sp. from Le Garouillas Oligocene locality, Quercy phosphorites, SW-France, is established in nomenclatural standards. 

A detailed comparative study of a complete skeleton of Bachitherium and a cladistic analysis of the sub-order Neoselenodontia lead us to propose a cladogram and a new classification of this group. The Tylopoda are the sister-group of the Ruminantia, which are chiefly defined by the fusion of the cuboid and navicular. Within this infra-order, Amphimeryx is the sister genus of a tetraselenodont group, in which the Hypertragulidae are well-separated group from a monophyletic group defined by the loss of trapezium, fusion of capitatum and trapezoid, and the isolation of the hypoconid on lower molars. The most primitive genera of this group, Lophiomeryx and Iberomeryx still have an open trigonid on the lower molars, but this is lingually closed in Archaeomeryx, sister-genus of the higher Ruminantia which have fused metatarsals and more evolved milk teeth. We divide them into two pan/orders : Tragulina (including the recent and miocene Tragulidae, and the North-American Leptomerycidae), and Pecora, with reduced lateral metacarpals and a new crest (telocristid) on the lower premolars. Within the Pecora, the upper molars of Gelocus are more primitive than those of Bachitherium (a genus with many autapomorphies in the dentition) itself more primitive than the group Prodremotherium + Eupecora, with fused metacarpals. We consider the Eupecora (including several genera without frontal appendages) to be monophyletic. 

Pangolins have never shown a high taxic diversity and their fossil record is scarce. We report here the first discovery of a partial humerus from late Oligocene deposits in Pech du Fraysse (MP28, France). The new specimen from Pech du Fraysse is described and compared to various extant and extinct species of pangolins. It shows a suite of morphological features very similar to the humeri discovered in Saint-André (MP 26), Solnhofen (Burdigalien), and Saulcet (Aquitanian), attributed here to Necromanis franconica. The description of the specimen from Pech du Fraysse allowed us to discuss the systematics of Paleogene and Neogene pholidotans. The differences between PFY 4051 and N. franconica on the one side, and N. quercyi on the other side, might be sufficiently important to justify a generic distinction. A comparison with extant species showed that N. franconica was likely terrestrial and fossorial based on its humeral morphology. 

The fossil mammals collected from the Eocene of Hammada du Dra (northwest Sahara. Algeria) and two fragmentary teeth from the Lutetian of M'Bodione Dadere (Senegal) are described.
The fossils from the northwest Sahara come from a lacustrian deposit dated by charophytes (Raskyella aff. pecki, Raskyella n. sp.. Maedleriella lavocati, Maedleriella sp. et ? Peckichara sp.) as Middle Eocene or perhaps Lower Eocene (Gevin, Feist and Mongereau, 1974). Several hyracoids (3 or 4) identified from this formation extends the age of the family Pliohyracidae Osborn in Africa. Three forms appear to belong in the genera Megalohyrax, Titanohyrax and perhaps Bunohyrax which have been know until now only from the lower Oligocene of the Fayum (M. gevini n. sp. ; T. mongereaui n. sp.. ? Bunohyrax or Megalohyrax indet.). Another hyracoidof small size is referred to a new genus, Microhyrax (M. lavocati n. sp.).
Helioseus insolitus n. g. n. sp. is described without ordinal assignment. Azibius (Sudre, 1975) which has been the subject of questions and interpretations is reviewed.
Only one tooth from the Lutetian of M'Bodione Dadere is complete enough to interpret. lt probably belongs to a condylarth and demonstrates for the first time, the presence of the order in Africa. The second tooth is too fragmentary for comment.
In conclusion., the paleobiogeographic role of Africa at the end of the cretaceous and the beginning of the Cenozoic is discussed. 

Six orders of birds adapted to aquatic life are represented among the numerous avifauna of "Saint-Gérand-le-Puy": Gaviiformes, Procellariiformes, Pelecaniformes, Ciconiiformes, Phoenicopteriformes, and Anseriformes. The present study of this avifauna proposes several changes in systematics:- Procellariiformes: Puffinus arvernensis does not belong in Procellariidae but in Diomodeidae, and it is transferred to the fossil genus Plotornis previously described in the Middle Miocene of France. - Pelecaniformes: Phalacrocorax littoralis remains in Phalacrocoracidae; P. míocaenus is different from the modern species, and is transferred to the new genus Nectornis. Empheresula arvernensis, described in the Oligocene deposits of Gannat, seems to be present in Saint-Gérand-le-Puy too. Pelecanus gracilis shows many differences from the modern species, and belongs to the new genus Miopelecanus, - Ciconiiformes: Ardea formosa nom. oblit. is a synonym of Proardeola walkeri. - Anseriformes: a new species closely related to swans is described, and belongs to the fossil genus Cygnopterus, of the Middle Oligocene of Europe; this species is called C. alphonsi. The ecology of each species is suggested by comparison with that of its nearest living relatives, and by study of osteological adaptations. 

The study of endocasts, or casts of the endocranial space, have played an important role in shaping understanding of mammalian, and particularly primate, brain evolution. Recently, the reconstructions of three-dimensional virtual endocasts from high-resolution computed tomography images have allowed for the visualization and quantification of endocasts in several Paleocene and Eocene primate species. Here we present the virtual endocast of MaPhQ 289 (informally known as the Montauban 9 skull), a specimen of Necrolemur antiquus Filhol 1873, a middle to late Eocene European primate of the family Microchoeridae. The virtual endocast of MaPhQ 289 reveals a lissencephalic surface morphology with expanded temporal poles and minimal overlap of the cerebellum or olfactory bulb by the cerebrum, which closely resembles the morphology of the endocast of its contemporary relative, Microchoerus erinaceus (Primates, Microchoeridae). MaPhQ 289 yields an endocranial volume (ECV) of 2.36 cm3, about 60% smaller than the volume of the most commonly cited ECV of N. antiquus. Thus, the size of the brain of N. antiquus relative to its body size is likely to be smaller than has been reported in previous literature, highlighting the importance of corroborating older ECV estimates with new evidence using 3-D imaging techniques. 

The first lists of Insectivores (Erinaceidae, Talpidae and Soricidae) from the Pliocene beds of Southern France and North-East Spain are given in this paper. The material from twelve localities is studied. These localities are geographically situated in Languedoc (Celleneuve, Vendargues, Nîmes, Sète, Balaruc 2 and Seynes), in Roussillon (Terrats, Serrats-d'en-Vacquer, Château d'eau and Mont-Hélène) and in North-East Spain (Layna, Medas Islands and Puebla de Valverde). These faunas correspond to the Early, Middle and Late Pliocene. 1 to 8 taxa are identified in these localities and 14 specific taxa are presently listed for this period in this area. Two new specific taxa are described as Galerix depereti nov. sp. from all the Early Pliocene localities in the North-Pyrenean area and as Desmanella gardiolensis nov. sp. from Balaruc 2. For this small mammals, two faunal assemblages are recognized. The first one is dated from the Early Pliocene (F 1, 2 and 3 zones in Aguilar et Michaux) and is characterized by Galerix depereti and rare and little diversified Soricids. The second one is Late Pliocene in age (zones G 2 and G 3). The fossils of the genus Talpa are relatively abundant and the Soricids are diversified and very abundant. The Middle Pliocene (zone G 1) is a transitional period. ln these faunas, most of the insectivore genera are known from the European Late Miocene beds (8 on 10). This fact demonstrates a relative continuity between the invectivore faunas from the Late Miocene to the Early Pliocene. In conclusion, somme paleoecological considerations are suggested.
  

Biographie et liste des publications de S. Garimond. 

 Detailed anatomical description of arsinoithere cranial remains from the Lower Oligocene, Fayum Depression, Egypt, provides the basic data for a systematic investigation. All cranial and some postcranial features are assessed from a phylogenetic standpoint. Several soft tissue characters are then added to a cladistic analysis based on 54 derived ungulate morphological characters. The resulting phylogenetic hypothesis implies that perissodactyls, sirenians, proboscideans and arsinoitheres constitute a monophyletic unit (5 synapomorphies). However, increasing the tree length by 3 steps reveals a closer association between hyraxes and perissodactyls. Nevertheless, 13 synapomorphies link proboscideans, sirenians and  arsinoitheres to the exclusion of all other ungulates. Form of the sphenopalatine and ethmoid foramina, recurved posttympanic process, absence of a fenestra rotundum in the petrosal, vestigial paroccipital process of the exoccipital and the highly unusual absence of a hypoglossal foramen in the skull, imply a robust sister-group relationship between arsinoitheres and proboscideans. In this analysis artiodactyls share only one derived character with all other ungulates studied. Monophyly of Ungulata, including Artiodactyla, is therefore only weakly supported. It is argued that pedal anatomy of hyraxes is non-homologous with that of Tethytheria. Arsinoitherium should now be classified within Tethytheria, sharing a sister-group relationship with Proboscidea. Hyraxes are excluded, thus refuting the concept of Paenungulata. However, monophyly of the wider concept, Pantomesaxonia, containing hyraxes, perissodactyls, sirenians, proboscideans and now, arsinoitheres, is supported by this study.