Les Pistes de Vertébrés du Stormberg Supérieur ("Trias terminal à Rhétien"), ou Quthingien

Si les zones du Stormberg inférieur se sont révélées contenir de nombreuses traces, surtout dans les faciès dits "Molteno moyen et supérieur", représentant apparemment la base du Keuper, il est frappant de voir pratiquement l'ensemble de cette grosse faune "Molteno" disparaître avec la fin de cette période, que nous avons appelée le "Maphutsengien".

Dès les premières zones du Stormberg supérieur, que nous nommons le"Quthingien" la zoocénose et la phytocénose, en même temps que les données d'ensemble manifestées par l'environnement, sont modifiées. Nous ne verrons plus guère de dépôts marécageux à flore riche et variée, parfois même luxuriante. Les fougères elles-mêmes ont disparu. Elles sont remplacées par de maigres plantes, aux feuilles très souvent filiformes qui paraissent témoigner d'un climat continental. Le sol est devenu de plus en plus rouge, avec des variations latérales beaucoup plus accusées. Les fleuves amenant des galets des monts du "Grand Sud" ont tari. La faune va en subir les conséquences. Certaines des espèces se révèleront sautillantes ou coureuses, pour un grand nombre plus légères et pour la quasi-totalité d'apparence carnivore ou entomophages, les phytophages devant se contenter d'un régime ingrat,difficile ou à tout le moins irrégulier,les dépôts le montrent.

C'est dans ces conditions que s'inaugure notre Etage nouveau,quelque peu discordant sur les zones A/5, A/6 ou A/7 du Stormberg inférieur (Maphutsengien). Le Stormberg supérieur (ou Quthingien) commence avec le paléopaysage remarquable dit de Moyeni, que nous allons maintenant étudier, typologiquement, avec ses homologues du même âge. Quelques 38 types d'animaux tous nouveaux vont défiler à nos yeux lors de la zone de base de cet Etage, ou zone B/1.

L'on nous avait proposé d'intituler ce Ile Tome de la série : "La grande Dalle de Moyeni et ses homologues. Paléo-spectacles, scènes et paysages animaux au Lesotho à l'approche du Trias finissant". Nous avons préféré garder le sous-titre plus haut, peut-être plus prosaïque.

Un llle Tome est en préparation : "Les développements ultérieurs et terminaux de la faune du Gondwana".

  

Extensive collecting in previously known and new Middle Eocene rodent localities in the Subathu Group of the Rajauri and Reasi districts, Jammu and Kashmir (northwest Himalaya, India) has yielded over 500 isolated cheek teeth, numerous incisors and cheek tooth fragments, and two mandibular fragments with M/1-M/3. An analysis of this additional material and a restudy of that reported earlier from the Metka (Rajauri) area has revealed the presence of diverse Middle Eocene ctenodactyloid rodents in India. The assemblage comprises six genera and twelve species, viz.., Bírbalomys woodi, B. ibrahimshahi, B. sondaari, Basalomys vandermeuleni, B. ijlsti, B. lavocati, Chapattimys wilsoni, C. debruijni, Gumbatomys asifi, cf. Advenimus bohlini, cf. Petrokoslovia sp. indet. 1 and cf. Petrokoslovia sp. indet. 2. Most species of this assemblage are endemic to the Indian subcontinent and are referred to the Family Chapattimyidae. However, at least three taxa, viz.., cf. Advenimus bohlini, cf. Petrokoslovia sp.indet. 1 and cf. Pezrokoslovia sp. indet. 2 show close affinities with their Central and Eastern Asiatic contemporaries and are attributed to the Family Yuomyidae. The dentitions of most of the Middle Eocene rodents from India are described here in better detail thus providing more precise characterization than was available earlier. The taxonomie status of Basalomys, formerly a subgenus of Birbalomys has been raised to that of a genus. Basalomys vandermeuleni is here designated as the type species of Basalomys in place of B. ijlsti. The rodent assemblage from the Subathu Group is specifically similar to that from the coeval Kuldana Formation of Pakistan. The absence of Birbalomys ibrahimshahi and cf. Advenimus bohlini in Pakistan is related to sampling bias. The identifications of the Early Eocene rodents from Barbara Banda, Pakistan have been reexamined and the familial assignments of some of the Early and Middle Eocene rodents from Central and Eastern Asia have been discussed and reviewed in the light of recent discovery of early rodents in Mongolia.

  

The Middle Eocene Fossillägerstätte of the Geiseltal lignite beds near Halle/S. (German Democratic Republic) is the reference locality of the European land mammal age Geiseltalian and of the Mammalian Paleogene reference levels MP 11 - MP 13. Due to this importance a reinvestigation is given of the lithostratigraphical development of the Geiseltal beds and of their vertebrate sites. The last are genetically related to the southwest border of the Geiseltal depression and the influx of carbonate-rich waters. The geographical distribution and stratigraphical position of the fossiliferous sites depends on subrosive and tectonically controlled distribution of coal seams. The geological factors and the known stratigraphical guide of some mammalian species suggest corrections of the age of some sites. Four of the alltogether five coal bearing phases contain the 35 sites with mammalian remains. By the distribution of the around 69 mammal species are characterized, with 5 faunal steps ranging from MP 11 to MP 14 or over the Geiseltalian up to the Lower Robiacian. Well distant are the faunas of MP 11 and MP 12. Beginning with MP 12 up to MP 13/14, the fossil record is very frequent by 27 sites. This evidence coincides somewhat more with the concept of land mammal ages compared to that of the punctual mammalian reference levels. 

The review of material recently collected in the new localities from the “Phosphorites du Quercy", and different localities from the South of France, bring new informations on the genus Eucricetodon THALER. 1966, and Pseudocricetodon  THALER,  I969 (Middle and Upper Oligocene. Western Europe). Thanks to Eucricetodon huerzeleri VIANEY-LIAUD, 1972, which were unsufficiently known until now, is proposed. During the middle Oligocene Eucricetodon atavus  MISONNE, 1957 seems to give rise to two lineages. One of them led to Eucricetodon huberi,which however exhibits a larger size and a development of progressive characters on the teeth. The other would be Eucricetodon huerzeleri well differentiated at the “Mas de Pauffié" standard level (beginning of the upper Oligocene). The ornementation of lower incisors is described, when possible. Though the fossils are not abundant, it seems that the ancestral lineage, Eucriretodon atavus, remains in  the upper Oligocene (Boningen standard level). evolving into Eucricetodon praecursor SCHAUB, 1925 (Rickenbach standard level). The characters of Eucricetodon dubius  (SCHAUB. 1925), represented by a numerous population in Pech Desse and Pech du Fraysse (Quercy). confirm that this species and Eucricetodon praecursor  belong to two different lineages. As Eucricetodon dubius shows more primitive features, this species could not originale from Eucricetodon atavus -Eucricetodon huberi. The appearance of this species at the level of Mas de Pauffié could be the result of an immigration. A new definition of Pseudocricetodon incertus (SCHLOSSER. 1884) is given. This species has been found in several localities, where it had not been identified until now. lts comparison with Pseudocricetodon moguntiacus  (BAHLO. l975), found at several localities from the standard level of Antoingt (end of middle Oligocene). shows a parallel evolution to that of Pseudocetodon incertus, which is  of larger size and with a less complicated pattern of teeth. 

 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. 

A skull of Parabos cordieri, undamaged by fossilisation, allows us to make an endocranial cast, which is compared to those of Bos taurus, Boselaphus tragocamelus and Hippotragus equinus. These are studied by general form (shape, volume, « rolling-up›› and inclination), and by detailed morphology (gyri and sulci patterns, rete mirabile, importance of rhinencephalon and cerebellum).
A preliminary study is made in order to recognize convolution topography, gyri, sulci, and the various encephalic regions as clearly on an endocranial cast as on a bare encephalon; an intermediate stage is the study of an encephalon covered with «soft» meninges.
The comparative study reveals great similarity between the endocranial cast of Parabos cordieri and that of Boselaphus tragocamelus, both belonging to the tribe Boselaphini. On the other hand, a marked differences separates the cast of Parabos cordieri from that of Bos taurus, representing the tribe Bovini to which Parabos cordieri had been referred. The endocranial cast of Hippotragus equinus, which had long been considered the descendant of Parabos cordieri, is also quite different from that of Parabos. 

Two species lineages of North American late Paleocene Plesiadapis exhibit a pattern of size divergence from a common ancestral lineage. Time series of fossils in each of these lineages are analyzed to test the idea that size divergence represents competitive character displacement. The critical factor in a test of character divergence is showing that divergent lineages evolved directionally rather than randomly (multifactorially). Analysis of evolutionary rates and their temporal scaling in Plesiadapis shows that both divergent species lineages have the scaling slope expected for lineages evolving randomly rather than directionally, and size divergence in Plesíadapis does not represent character displacement. Rates of evolution commonly observed on a per-generation time scale are high enough to produce character displacement within a few generations. Thus character displacement is not likely to be visible on scales of time that can be studied in the fossil record. 

Avant propos au "Premier catalogue des specimens-types Paléontologiques déposés dans les collections de l'Université de Montpellier II". 

The genus Microstonyx was found only in the north eastern part of Spain : M. antiquus, referable to the
Eppelsheim type, from the pyrenean basins ; M. antiquus, more progresive, from the Vallès Basin : M. major-erymanthius group, from the Teruel-Ademuz Area and from Pieru (Penedès Basin).

  

La poche de Ste-Néboule, commune de Béduer (Lot), 15 km environ à l'WSW de Figeac, fait partie du groupe le plus septentrional des gouffres creusés par les ruissellements du Paléogène dans les calcaires jurassiques de la bordure sud-ouest du Massif Central et qui furent comblés à la même époque par des argiles sidérolithiques accompagnées de phosphate de chaux concrétionné ainsi que des restes de la célèbre faune dite «des phosphorites du Quercy» . 

A new artiodactyl, Diacodexis antunesi n.sp., is described from the early Eocene of Silveirinha, Portugal. Comparisons are made with Diacodexis gazini GODINOT, 1978, D. varleti SUDRE et al., 1983, D. cf. varleti from Paris Basin sites, D. sp. from Dormaal and from localities in Spain and England, D. secans from North America and D. pakistanensis from Asia; affinities and evolutive tendencies are discussed. The presence of Diacodexis in the locality of Silveirinha confirms the very early Eocene age of the latter. As Diacodexis antunesi appears ta be more primitive than D. gazini from Rians (early Eocene of France), it lends corroboration to the interpretation (essentially based previously on condylarths) of the Silveirinha assemblage as the oldest Eocene fauna known in Europe and supports the hypothesis that early artiodactyls migrated from Europe to North America. 

The Caudata are known by two Salamandridae ; one of them is attributed to the genus Megalotriton. The Pelobatidae form the major part of the Anura ; a few bones indicate also the presence of Neobatrachia. 

Les Carnivores Fissipèdes de Sainte-Néboule appartiennent tous au genre Cynodictis et semblent constituer une population homogène. Celle-ci se distingue suffisamment des espèces déjà décrites pour constituer un taxon particulier : Cynodictis lacustris neboulensis n. s. sp. . L'étude des variations à l'intérieur de cette population nous a conduit à reconsidérer les critères utilisés pour définir les espèces existantes et à regrouper certaines d'entre elles. Il semble qu'il demeure cependant trois lignées distinctes dans le genre Cynodictis mais le matériel nous paraît encore insuffisant pour traduire cette remarque en termes de systématique. 

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.

  

The reproductive biology of living birds differs dramatically from that of other extant vertebrates. Distinctive features common to most birds include a single ovary and oviduct, production of one egg at daily or greater intervals, incubation by brooding and extensive parental care. The prevalence of male parental care is most exceptional among living amniotes. A variety of hypotheses exist to explain the origin of avian reproduction. Central to these models are proposed transitions from a condition of no care to maternal, paternal or biparental care systems. These evolutionary models incorporate a number of features potentially preservable or inferable from the fossil record (integument, skeletal adaptations for flight, egg and clutch size, nest form, hatchling developmental stage, the number and function of oviducts, and the mode of egg incubation). Increasing availability of data on dinosaur reproduction provides a means of assessing these hypotheses with fossil evidence. We compare dinosaur data to a selection of models that emphasize maternal, paternal or biparental care. Despite some congruence with dinosaur features, no single model on the evolution of avian reproduction conforms fully to the fossil record, and the ancestral parental care system of birds remains ambiguous. Further investigation into dinosaur parental care, nest structures, clutch geometry, egg-pairing, eggshell porosity, and embryo identification may eventually resolve these issues.  

This paper is mainly devoted to the systematics of rodents from two middle Eocene (Bartonian) localities: Chéry-Chartreuve and Rocourt-Saint-Martin (Aisne, Eastern Paris Basin). These two localities are stratigraphically located slightly above the Auversian sands. The two faunas, which comprise 11 and 8 taxa, respectively, are very different in their composition. That of Rocourt-Saint-Martin shows strong similarities with that of the geographically very close locality of Grisolles, referred to the MP16 mammalian Reference level. The very distinct fauna of Chéry-Chartreuve includes a new species of Ailuravinae, Ailuravus nov.sp, and some teeth of the theridomyid Protadelomys, which represent archaic elements in the fauna. The most abundant species of the locality represents a new genus of primitive Theridomyidae. The presence of some teeth belonging to a new species of large Remyinae, Remys nov. sp., of Elfomys engesseri HOOKER & WEIDMANN, and a population of small dimensions referred to the genus Estellomys allow a correlation with Les Alleveys (Switzerland), with however some differences that would indicate an older age for Chéry-Chartreuve. Situated at the base of the "Marinesian" from the Bassin de Paris, this fauna is unquestionably different from those referred to the MP16 reference level and could represent an older level for which the macrofauna remains very poorly known. Conversely, the comparison of rodents from La Livinière II with those present in MP16 faunas, especially those of Robiac (Gard), shows a great similarity between both localities. This casts doubts on whether to keep this La Livinière II faunule to define the current MP15 reference level, while the biostratigraphical position of Pontils 26 (Spain), previously referred to this level, is reconsidered. Chery Chartreuse could be a good candidate for a new definition of the MP15 reference level. 

Description, for the first time, of the skull of Ruscinomys Depéret on the basis of a nearly complete specimen, and description of a new facial part of a Megacricetodon Fahlbusch skull (material from upper Miocene, Spain). New description of the skull (facial part) of " Cricetodon" incertum Schlosser on the basis of the specimen from the Oligocene of Quercy phosphorites already published by S. Schaub.
Comparison of the skull of “ C. ” incertum with that of the asiatic genus Cricetops Matthew and Granger and that of the North-American Eumys Leidy. These three genera of similar age display very distinct characteristics; their common origin must go back to the Eocene, as Wood believed.
Comparisons of the auditory regions of Ruscínomys and of Megacricetodon where important differences are noted. Comparison of the Cricetodontines from European Miocene with contemporary Cricetines. Contrary of the opinion held by Schaub, their anatomical differences do not seem suflicient to warrant the contention that none of the Cricetines derive from a form close to one of the known Cricetodontines. 

Perutherium altiplanerise THALER, 1967 from the Late Cretaceous of Peru has long been recognized as South America's oldest known placental mammal. Since its description Perutherium has been generally regarded as having condylarth affinity Based on our identification  of a unique notoungulate synapomorphy we recognize Perutherium as the oldest and the most generalized known member of that order. This new determination and the large taxonomic diversity (five families) of notoungulates in rocks of Paleocene age in Argentina and Brazil, favor a South American origin for this group. The occurrence of notoungulates in rocks of Late Paleocene age in Asia and North America is explained by dispersal of a notoungulate stock from South America to North America and from there to Asia. 

The study of the bovids from Al Jadidah (Hofuf Formation, Saudi Arabia) confirms that the fauna comes from a pre-Hipparion level. The Al Jadidah age is close to that of Fort Ternan (14 m.y.) and Beni Mellal, but cannot be older than that of Fort Ternan. The age of the Hofuf Formation is close to but slightly older than the oldest deposits of the Ngorora Formation (Kenya). 7 to 9 species have been recorded, of which 2 to 4 remain indeterminate. If the great specific diversity of te bovids from this locality gives evidence of immigrations from anterior Asia (Turkey) (e.g. Pachytragus Iigabuei sp. nov.), the bovid assemblage of Al Jadidah results in fact from a double influence: from the anterior Asia and mainly from Africa (e.g. the Caprotragoides lineage and the Neotragini? Homoiodorcas). The Al Jadidah bovids reflect, on the whole, the predominant character of open to very open environment, which supports the conclusions drawn from our two preliminary studies. 

On the Track of Ice Age Mammals justifie pleinement son titre car l'auteur, Anthony J. Sutcliffe, apporte au lecteur faits et interprétations qui l'amèneront à s'intéresser encore plus au passé récent et à l'avenir de son environnement et à la question de l'impact de l'homme sur la nature. Après les chapitres qui présentent les temps glaciaires et les divers témoignages qui nous en sont parvenus, les cinquième et sixième apportent les informations nécessaires à la compréhension des résultats que nous donnent les chercheurs: principes, moyens d'étude et limites des méthodes, difficultés de l'intégration des données dans un cadre stratigraphique, variabilité des signaux climatiques, variabilité de leur intensité selon l'endroit par rapport au centre de la glaciation, complications liées à la qualité inégale de l'enregistrement géologique, en mer et sur le continent.