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

A 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 utilization of screen-washing and attack by dilute acetic acid has permitted the collecting, in the Miocene of the department of Hérault (France), of a very rich ichthyofauna. This fauna is presently comprised of about 60 studied species, of which 11 are new, and represents, in the present state of knowledge, the most varied Miocene selachian fauna described in the world.
The abondance of material has allowed an overall revision to be made; it has thus been possible to complete the description and the figuration of species that were poorly known until now, and to synonymize species that were established on simple morphotypes. Paleo-ecologic study of the ichthyofauna has permitted conclusions to be drawn relative to climate and bathymetry; it was thus possible to show that the Miocene fauna of Hérault was a fauna of a subtropical sea, essentially neritic with rare pelagic contributions.
Knowing the stratigraphic position of the localities, it has been possible to distinguish three faunal assemblages based on associations of species. Some hypotheses on the evolution of certain lineages have been expressed.
The comparison of this fauna with that of other regions permitted the relationships of two diflerent faunal provinces to be specified: the first belongs to the northern domain, characterized by a fauna still subtropical but with numerous temperate water elements; the leoond belongs to the Mesogean domain characterized by warm water forms. It has also lhovm that contemporary faunas could be very different according to the bathymetric zone in which they lived, which furnishes valuable information for the paleogeographic reconstruction of sedimentary basins.
  

The genus Cryptomeryx SCHLOSSER, 1886, inusited during a long period, has been discovered in Lower and Middle Oligocene localities of the Quercy region (South-West France). This new material, as well as specimens from the old collections referred to Cryptomeryx, are described; their study, allows us precising the definition of the genus, and confirming its allocation to the Tragulidae family. The type species of the genus, Crypmmeryx gaudryi (= Lophiomeryx gaudryi FILHOL, 1877), occurs in several localities at the base of the Middle Oligocene (Itardies, La Plante 2, Roqueprune 2, Soulce, Herrlingen 1). The new species C. matsoui n. sp. has been defined in the older locality of Mas de Got (top of Lower Oligocene). It is possible that the species Pseudamphimeryx decedens STEHLIN, 1910 pertains to the same genus. Also to the Tragulids must be referred the monospecific genus Iberomeryx (I. parvus GABOUNIA, 1964) from Upper Oligocene of Benara (Georgie, URSS), with which Cryptomeryx is related. These genera are not direct ancestors of Miocene tragulids; their occurrence in the Western European Oligocene results from a first immigration wawe of the family. These Tragulids are one of the most archaic groups of Ruminants. They are probably derived from a primitive stock which had acquired in Asia the selenodont condition of the dentition. 

A fossil fish fauna, based on 5533 otoliths, from the La Paillade locality at Montpellier is described and figured. The otolith-bearing marls correlate to mammal zone MN l (Aguilar, 1982), and thus represent the earliest Miocene. The fish fauna consists of 30 taxa belonging to 20 families. Two species are new: Dussumieria sittigi and Liza gaudanti. The predominant faunal element is the Lesueurigobius vicínalis-species complex, composing 73% of all investigated otoliths. The palaeoecological analysis reveals a marine to euryhaline fish fauna living under tropical to subtropical conditions in the transition zone littoral - sublittoral. Water depth probably was more than 10 m. The scarcity of pelagic físhes suggests that the habitat was either a sheltered bay and/or far away from the open sea. Furthermore, some genera represented in the La Paillade fish fauna presently live exclusively in the Indopacific realm. Their presence strongly supports a broad connection between the Indian Ocean, the Mediterranean, and the Paratethys Seas during the Early Miocene (Aquitanian). From a palaeobiogeographical point of view, faunal relationships were found between the La Paillade fish fauna and both the Paratethys fish fauna and the fish fauna from the deposits in the Upper Rhinegraben and the Mayence and Hanau Basins (Germany).



  

The East African Miocene Macroscelidea, lnsectivora and Chiroptera are revised on the basis of new material. New taxa proposed are: Miorhynchocyon, .n. gen. (Macroscelididae): Míorhynchocyon meswae, n. sp.: Pronasílío ternanensis. n. gen.. n. sp. (Macroscelididae); Hiwegicyon juvenalis, n. gen. n. sp. (Macroscelididae); Parageogale, n. gen. (Tenrecidae): Prochrysochlorinae, n. subfam. (Chrysochloridae): Propottininae, n. subfam, (Pteropodidae); Chamtwaria pickfordi, n. gen., n. sp. (Vespertilionidae). Gymnurechnínus songhorensis is synonymised with G. camptolophus. The new material provides additional information on the dentition, especially of Myohyrax oswaldi. Galerix africanus. Amphechínus rusingensis, Protenrec tricuspis and Parageogale aletris. Partial skulls are described of Amphechinus rusingensis, Protenrec tricuspis, Prochrysochloris míocaenicus and Taphozous incognita. The oldest member of the Macroscelidinae (Pronasilio) is described from Fort Ternan. Galerix africanus is closely related to G. exilis from Europe. Amphechinus rusingenesis is compared with Asiatic Oligocene Erinaceinae. The Miocene age of Crocidura is rejected. On the evidence of humeri, the following families of Chiroptera are newly reported: Pteropodidae, Nycterididae, Vespertilionidae, Molossidae. Propotto is regarded as an offshoot from the Pteropodidae, not ancestral to modern forms. Chamtwaria is a primitive vespertilionoid, provisionally placed in the Kerivoulinae. Erinaceidae probably entered Africa at the beginning of the Miocene, before 20 Ma. Faunistic differences between deposits are largely to be ascribed to differences in local environment. 

In the course of the current revision of adapid primates from the Eocene Geiseltal, an almost complete specimen was found in the Geiseltal Museum collections. The fossil, the most complete adapid specimen so far discovered in Europe, has been determined as Pronycticebus neglectus n. sp.
Ecology and locomotion as well as the likely phylogenetic position within the infraorder Adapiformes are discussed. 

A cheek tooth recently unearthed in the Lopar Sandstone unit, of late Eocene age, in the northern part of Rab Island, Croatia, is one of the very few Eocene mammalian remains found in the Adriatic area. Thorough comparison of this tooth with those of Old-World Palaeogene mammalian orders suggests that it is a M3 belonging to an embrithopod afrothere. The specimen is referred to as Palaeoamasia sp. This genus was formerly known only in Eocene deposits of Anatolia but with close relatives in Romania among Palaeoamasiinae. The geographical distribution of this subfamily perfectly matches the recently-named Balkanatolian landmass, which experienced in-situ evolution of endemic mammals prior to the Grande Coupure event that occurred around the Eocene–Oligocene transition. This last event is characterised by massive Asian immigration in Western Europe and the supposed extinction of many endemic Central and Western European mammals, including Palaeoamasiinae.
  

Situated in the Cape region of the Republic of South Africa (RSA), the paleontological site of Langebaanweg is dated to 5.1 Myr and is famous for having yielded an abundant vertebrate assemblage, including numerous rodent species from the Mio-Pliocene transition. Based on molar morphology and skull anatomy, the single Gerbillinae taxon identified at Langebaanweg and described in this paper is allocated to Desmodillus, which is a modern monotypic South African endemic genus. It is significant in being the oldest representative of the genus in Africa. We describe here a new species of this genus which is larger than the modern D. auricularis, but nevertheless retains some of its main characteristics, namely the shape of the maxilla and mandible, the presence of poorly fused alternating cusps, and no longitudinal crest. This taxon differs from modern South African Gerbilliscus representatives in some mandibular and maxillary characters, in the m1 prelobe cusp, and in having less fused cusps. Two fossil Gerbillinae discovered in the Upper Miocene of Africa and Asia, Abudhabia and Protatera, have been compared with the new species. We discuss their relationships with modern and Plio-pleistocene Gerbillinae and conclude that Abudhabia could be the sister taxon of Desmodillus and that around 6-5 Myr a vicariance event allowed Gerbillinae to diversify into modern Desmodillus in South Africa, and Gerbilliscus in East Africa. The murine/gerbilline ratio, which is a good indicator of rainfall, supports other proxies which suggest that at 5.1 Myr the climate in the Langebaanweg region was more humid than today. 

The Tafna Basin corresponds to the lowlands, which are located in front of Tessala and Traras ranges, below the Tlemcen mountains, Algeria. This basin displays a complete sedimentary cycle dominated by lagoonal-fluvial and marine deposits. The continental formations located at the base of these deposits are mainly composed of alternating sandstones and clays. An early late Miocene age has been previously attributed to them, based on direct correlations with marine deposits. Search for micromammal fossils led to the discovery of three different rodent species from a single level of the Djebel Guetaf section, located at the bottom of these deposits. The rodent assemblage indicates a late Miocene age. Combined magnetostratigraphical and biostratigraphical investigations were carried out to provide a more accurate age control of these continental deposits. Sixty-four oriented rock samples were collected for a magnetostratigraphic study along a 92 meters thick section including the fossiliferous layer. Rock magnetic investigations indicate the presence of both high and low coercivity minerals. Specimens subjected to progressive thermal demagnetization procedures show that the samples exhibit a high temperature magnetization component and display a normal polarity. Based on biostratigraphic constraints, the Guetaf section is correlated with Chron C4An, indicating an age ranging from
9.1 to 8.7 Myr.
  

In the course of a revision of the Equoidea numerous dentitions as well as a partial skeleton of a Phenaeodont were discovered from the Middle Eocene lignite beds of the Geiseltal locality. These fossils are recognized as a new genus and species of Phenacodontidae : HaIlensia matthesi n.g. n.sp.. The species is present in the « untere und obere Unterkohle ›› (uUK, oUK = the lower and upper part of the Lower Coal Seam) as well as in the « obere Mittelkohle ›› (oMK = the upper part of the Middle Coal Seam). Two fragmentary upper jaws described and figured by Matthes (1977) as Propachynolophus gaudryi are also belonging to Hallensia matthesi. Thus the decisive argument for classifying the " Unterkohle " of the Geiseltal section as Lower Eocene has to be dropped. Another relict form of the Geiseltal is Esthonyx tardus n. sp. documented by a fragmentary mandible coming from the « untere Unterkohle ››. This is the latest Tillodont from Europe. Contrasting to E. munieri from the european Lower Eocene the dentition of E. tardus is morphologically more progressive. 

An ilium from the Upper Eocene (MP l7a) of Hordle, England, represents the first report of Rana from the Eocene of Britain. The ilium is similar to those of the water frog (Rana [ridibunda]) species group. 

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. 

Fourteen distinct phyletical lineages which belong at least in three families: Ischyromyidae ALSTON, 1876, Gliridae THOMAS, 1896 and Theridomyidae ALSTON, 1876, have been identified after the study of more than 3600 rodent dental remains from about twenty Early and Middle Eocene european localities. A systematical and phylogenetical revision of these rodents has been achieved. Nearly all the specific and generic diagnosis are emended. Several new combinations and synonymies are proposed. Four new species and two new genera, Euromys nov. (Ailuravinae) and Hartenbergeromys nov. (Microparamyini), are named and described. Euromys nov. gen. is known by three distinctive ypresian (MP 7 to MP 10 european reference levels) chronospecies. This new lineage is thought to be the direct ancestor of Meldimys MICHAUX, 1968 and Ailuravus RUTIMEYER, 1891. A new species of the genus Plesiarctomys BRAVARD, 1850, Pl. lapicidinarum from Condé-en-Brie (MP 8-9 reference level), allows to relate the Plesiarctomys lineage to the Pseudoparamys MICHAUX, 1964 one. The taxa Sparnacomys HARTENBERGER, 1971, Pantrogna HARTENBERGER, 1971, and Corbarimys MARANDAT, 1989 are erected to genus rank; the last one is not thought to be an Ischyromyidae. A new chronospecies of Pantrogna, P. marandati nov. sp. from the locality of Prémontré (MP 10 reference level), is described. This lineage is at the origin of two others, namely Masillamys TOBIEN, 1954, including M. mattaueri (HARTENBERGER, 1975) nov. comb. (MP 10 reference level), and Hartenbergeromys nov. gen., known from MP 10 (H. hautefeuillei nov. sp.) and MP 11 (H. parvus TOBIEN, 1954) reference levels. The phylogenetical position of Hartenbergeromys nov. gen., at the origin of the european family Theridomyidae, is discussed. The systematical and phylogenetical status of two probable Paramyinae, "Paramys" woodi MICHAUX, 1964 and an unnamed genus and species, are discussed. New populations of the primitive Gliridae Eogliravus HARTENBERGER, 1971 and of the primitive Theridomyidae Protadelomys HARTENBERGER, 1968, are described and assigned to previously known species.

  

Le Quercy est aujourd'hui un vaste plateau calcaire, parcouru par un réseau karstique actif, pro· fondément 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.
Le Causse du Quercy devait être un plateau très disséqué par la karstification, à surface lapiazée creusée de gouffres en rapide évolution interne, et couvert d'un sol assez épais. Une forêt tropicale humide, avec notamment des Myricacées et des gymnospermes, recouvrait l'ensemble du pays, à peine interrompue au niveau des rares points d'eaux situés dans les bas-fonds. Dans ce cadre vivait une faune considérablement différente de la maigre faune actuelle. Si nous pouvons l'imaginer, c'est grâce aux cadavres des animaux entraînés dans les cavités par les eaux courantes ou les prédateurs, ou bien logeant et mourant dans les milieux souterrains. Nombre de ces restes, fossilisés, sont parvenus jusqu'à nous et sont aujourd'hui l'objet d'étude. 

The two species of Sandgrouse from Quercy, Pterocles validus MILNE-EDWARDS and P. larvatus MILNE-EDWARDS, are ascribed to the genus Archaeoganga MOURER-CHAUVIRÉ which includes a third, very large species, A. pinguis. The Sandgrouse of Saint-Gérand-le-Puy, Pterocles sepultus MILNE-EDWARDS, is ascribed to a new genus, Leptoganga. This form appears in the Upper Oligocene of Quercy, in Pech Desse and Pech du Fraysse localities, and is still present in the Lower Miocene of Saint-Gérand-le-Puy and Paulhiac. Recent Sandgrouse live in semidesert or desert areas. The indications provided by mammal and bird faunas in the localities where sandgrouse were found, confirm that the paleoenvironment was open and arid. The morphological study of these fossils indicates that, in the Upper Eocene, the Pteroclidae were already completely individualized with respect to Charadriiformes. 

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

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). 

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.