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".

  

A new Devonian vertebrste locality has been discovered in 1976 in the basal part of the Khush-Yeilagh Formation in the eastern Alborz Mountains of Iran. The fossils occur in a band one centimeter thick which is identifiable at other outcrops in the area. A preliminary study of the remains has yielded the following faunal list : Placodermata (Phlyctaeniida indet.. Groenlandaspididae indet.. Coccosteidae indet., Holonema sp., Ptyctodontida indet., Antiarcha indet., Bothriolepis cf. kwangtungensis), Elasmobranchii (Ctenacanthidae indet.), Acanthodii (Gyracanthus sp., « Onchus » overathensis, Ischnacanthiforme indet.), Dipnoi (? Dipteridae indet.), « Crossopterygii ›› (Onychodus cf. sigmoides, Holoptychiidae indet., Osteolepididae indet.). This fauna is older than the Middle Givetian and probably of Middle or Lower Eifelian age. Owing to its diversity, it may prove possible to use it in the future as a reference fauna for the study of the Devonian vertebrates in Central Asia and the Middle East. 

The rodents (Cricetidae, Gliridae, Sciuridae) found in lacustrine, brackish marine and karstic sediments of Miocene age in Languedoc, assign the position of the different localities in the scale of "niveaux repères" used by mammalogists. Some detailed stratigraphical studies bring several correlations between this continental biochronological scale and the marine scale ; the most important results are the Aquitanian age of the "niveaux repères" of Coderet and Paulhiac, the Burdigalian age of Laugnac, Estrepouy, Vieux-Collonges, La Romieu and Sansan and the Langhian or Lower Serravallian age of La Grive M. The correlations between the Tethys and the Central Paratethys for the Lower Neogene profit also of these results, since the locality of Neudorf Spalte 1, 2 (Czechoslovakia) is shown to be younger than Sansan (France). The paleontological study has also several geological inferences for the Miocene of Languedoc ; with the calibration of this Miocene, we know quite precisely that the Lower Miocene is chiefly a time lacustrine sedimentation, and also that the marine Miocene sedimentation ends early in the Miocene Period, in Langhian or lower Serravallian times. 

The anatomy of Norselaspis glacialis n.g., n.sp., a primitive kiaeraspidian from the Lower Devonian of Spitsbergen, is described on the basis of spécimens studied by grinding sections or prepared with dilute formic acid. This study yielded some new anatomical details, including the presence of a canal prolonging posteromedially the canal alloted to the facial nerve by Stensiö. This posterior prolongation of the « facial canal ›› into the posterolateral part of the labyrinth cavity is consistent with the hypothesis put forward by Allis, Lindström, Jefferies and Whiting, that this canal housed the glossopharyngeus nerve. Furthermore, in N. glacialis, the foramen usually referred to as the foramen for the œsophagus opens posteriorly into a cavity in the postbranchial wall, referred to here as the intramural cavity, and which is interpreted as having housed the heart. Consequently, the œsophagus probably accompanied the dorsal aorta through the aortic canal. Finally, the foramen generally interpreted as having transmitted the ventral afferent arterial trunk is here considered as having housed the hepatic vein, which emptied into the venous sinus of the heart. The ventral afferent arterial trunk may thus have passed through the former «œsophageal ›› foramen.
The problem of the position of the dorsal nerves in the Osteostraci is discussed, and it is suggested that the three foremost nerve canals opening into the oralobranchial cavity housed the maxillary ramus of the trigeminus, the facial nerve and the glossopharyngeus nerve respectively. The mandibular ramus of the trigeminus must have accompanied one of the two foremost nerves, but for the moment it is impossible to decide which.
The problem of the nature of the interbranchial crests of the Osteostraci is briefly discussed. Comparison with the branchial apparatus of the Petromyzontida does not support the hypothesis that the interbranchial crests are part of the branchial arches, incorporated into the endoskeletal shield. A different hypothesis is proposed, that the branchial skeleton of the Osteostraci was situated entirely inside the oralobranchial cavity, and was attached to the endoskeletal shield only by the ventromedial processes. The grooves classically allotted to the efferent branchial arteries would thus have housed extrabranchial arteries, branching off from the dorsal aorta, and irrigating the ventral branchial musculature.
A phylogeny and a classification of the kiaeraspidians are proposed. The evolution of this monophyletic group is characterized by, e.g., reduction of cornual processes, shortening of the abdominal division of the shield, subdivision of the lateral fields, and enlargement of the supraoral fossae.
The phylogenetic position of the kiaeraspidians within the Osteostraci remains uncertain. Their sister-group may be either the benneviaspidiens or the thyestidians, or Thyestes alone (in which case they would have to be included within the thyestidians). 

Messiphoca mauretanica nov. gen., nov. sp. représente le premier Phocidae fossile recensé en Afrique du Nord. Provenant du gisement sahélien (Miocène supérieur) de Raz-el-Aïn (Algérie), il est connu par quelques os du membre antérieur (humérus, radius, ulna), quelques vertèbres dorsales et un crâne très fragmentaire. La description de cette forme nouvelle amène à considérer Messiphoca mauretanica comme un Monachinae archaïque proche de l'origine du groupe Pliophoca - Monachus. L'incidence de cette interprétation d'un Phocidae «pré-Messinien ›› sur la crise de salinité du Messinien est aussi envisagée. 

The recent excavation of the Upper Miocene locality of Montredon (Hérault) provides a rich material of Mammals. The present study deals with rodents. The new quarry is 10 meters above the classical Depéret's Quarry. No differences have been noticed between the rodent populations coming grom these two sites. The description of two new species : Hispanomys mediterraneus and Muscardinus heintzi is given and the dormouse Eliomys is recorded there for the first time. The new zonation of the Miocene given by Aguilar (1982) and the previously established correlations between marine and continental chronological scales give Tortonien age (estimated between 9 and 10 M.Y. old) to the site of Montredon, which belongs to the Mammal zone D 2. 

This paper presents a preliminary list of insectivores from the Vallesian beds at Montredon (France). The associated rodent fauna has established a Vallesian age for the fauna. Eleven species belonging to the Soricidae, Talpidae, Erinaceidae, and Dimylidae are identified of which four only are referred with certainty to forms already named. 

The Montredon local fauna yielded very rare bats, represented by damaged isolated teeth. Only a few documents are available for this period of the European Neogene. ln this poor state of knowledge, the material represents three undetermined species, a supposed molossid and two vespertilionids. 

The Condylarths from Hainin (Hainault, Belgium) show no affinity at the generic level to those known in other Paleocene localities of Europe and North America ; they are described as new forms : Monshyus praevius n. gen., n. sp. and Prolatidens waudruae n. gen., n. sp. Monshyus praevius, discovered in only one of the levels in the excavation at Hainin, is similar to the genera Microhyus TEILHARD and Louisina RUSSELL ; with them it is included in the subfamily Louisininae (Hyopsodontidae). With respect to Microhyus and Louisina, Monshyus is distinguished by the precociously modern aspect of its upper molars, the only teeth that are referable. Prolatidens waudruae, known only by lower molars, was found in several levels in the pit at Hainin. It is an arctocyonid presenting possible relationships to the North American form Oxyprimus galadrielae ; it therefore has been provisionally attributed to the subfamily Oxyclaeninae. If this attribution is confirmed, this species will constitute the first and only representative of the group in Europe. 

The oldest european marsupials are described from some specimens (isolated upper molars) recently found from the Hainin sediment (Middle Paleocene of Belgium). These fossils document a new species of the Peradectes genus. They give evidence of a much older occurrence of the marsupials in Europe than it was assumed. They allow us to postulate a didelphid dispersal from South America towards the western-holarctic area operating in two phases : the first one of the Peradectes genus at the end of the Cretaceous; the second one of the Didelphíni tribe at the end of the Paleocene. A central american crossing is likely for the first one,  whereas a transafrican way is tentatively argued for the second one. 

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 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. 

Re-evaluation of the small collection of mammal fossils from the Changxindian Formation near Beijing, China indicates the following taxa are present: Eutheria, Hypsimilus beifingensis, cf. Miacis sp., Anthracotheriidae and Forstercooperia grandis. The presence of Forstercooperia grandis indicates an Irdinmanhan age and does not support previous assignment of a Sharamurunian age to the Changxindian Formation. 

Le volume 11 du «Handbuch der Paläeoherpetologie», consacré aux «Serpellles» et redigé par Jean-Claude Rage, apporte une remarquable information sur ce groupe particulier de reptiles dont on ne pouvait guere se faire une idée très précise à partir, par exemple, des seuls chapitres des traités classiques de Paléontologie et de Zoologie. 

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. 

The classifications of the recent vespertilionine bats were made wihtout taking in account the teeth morphology; this resulted in a reduction of the possibilities of comparison with the available fossils. The generalized use of dental formulae was abusive: this contributed to the admission of artificial genera. These conditions have long delayed the consideration of characters able to frame the phylogeny of the sub-family. In the first part of the study, the teeth morphologies are described and analysed. morphological reference types are established for each upper and lower tooth: they should make an easier elaboration of criteria for the differentiation at generic level. The position of the species in view of these criteria allows one to group them into homogeneous genera, and to appreciate the degree of relationship that the latter have between them. The second part of the study (next publicationà will develop inferences dealing with systematics and phylogeny 

The genus Parasteiromys AMEGHINO, 1904 is revalidated, and P. friantae sp. nov. (Hystricognathi, Erethizontidae) from Colhuehuapian (early Miocene) sediments of the southern cliff of Colhue-Huapi Lake (Province of Chubut, Argentina), is described. The molar morphology of these taxa and of living porcupines adds new elements to understand the dental evolution of the Erethizontidae, and to propose the hypothetical ancestral molar pattern for this family. This pattern does not correspond to any of the morphologies traditionally proposed as ancestral for South American hystricognathous rodents. The proposed pattern is characterized by a metaloph disconnected from the posteroloph and oriented towards the hypocone, and the third loph incompletely developed with the lingual portion homologous to the mesolophule of Baluchimyinae (Chapattimyidae) from the Miocene of Pakistan. The inferred steps of the molar evolution of erethizontids towards the pentalophodont condition, considered derived for the family, are illustrated. This study strengthens the hypothesis placing erethizontids in a basal position among rodents of the suborder Hystricognathi.

  

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 systematic analysis of an hipparionine horse assemblage from Maragheh, Iran is made. A brief orientation to systematic philosophy and informal superspecific characterizations of some Old World hipparionines is given as a background to this work. A character state analysis of skulls is made, and has revealed five distinct species. A character state and stratigraphic trend analysis of isolated check tooth and postcranial remains, with known provenance, is also made. These two combined analyses reveal that the most resolute discrimination of hipparionine species and their evolutionary relationships occurs when multiple character complexes of associated skulls, maxillary and mandibular dentitions are made. When this is not possible, skulls have provided the best basis for discriminating species and their evolutionary relationships. Traditional characters of isolated cheek teeth and postcranial remains are shown here to offer limited information content for hipparionine phylogenetic systematics. The systematic portion of this study includes a comprehensive description of cranial and postcranial remains, and has further corroborated the distinction of five species which belong to at least three superspecific groups including: «Hipparion» geltyi sp. nov., Group 1; Hipparion prostylum (s. l.), and Hipparion campbelli sp. nov., Group 3; «Hipparíon» aff. moldavicum and «Hipparion» ?matthewi, Group 2. These species stratigraphic ranges and evolutionary relationships are also given here and argued to be important for establishing future hipparionine geochronologic correlations between a number of Eurasian late Miocene provinces. 

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.