Remains of sea turtles from the Ikovo locality (Lugansk Region, Ukraine; Middle Eocene)

Fragmentary remains of sea turtles (Cheloniidae sensu lato: Argillochelys sp., Puppigerus nessovi Averianov, 2005, and Cheloniidae gen. indet.) from the Ikovo locality (Lugansk Region, Ukraine; Lower Lutetian, Middle Eocene) are described. The genera Argillochelys and Puppigerus are recorded for the first time in Eastern Europe. The turtle assemblage from Ikovo is similar at the generic level to West European assemblages (Belgium, Great Britain) of approximately the same age (Lower-Middle Eocene). In the presence of P. nessovi, the Ikovo assemblage is similar to that from the Middle Eocene Dzheroi 2 locality (Uzbekistan).     

On a tomistomine crocodile (Crocodylidae,Tomistominae) from the Middle Eocene of Ukraine

A dentary fragment and osteoderm of the crocodile Tomistominae indet. from the Middle Eocene (Early Lutetian) Ikovo locality (Lugansk Region, Ukraine) is described.     

Middle Eocene Pelagornithidae and Gaviiformes (Aves) from the Ukrainian Paratethys

Abstract: We describe avian remains from Novopskov, a new middle Eocene marine locality in Ukraine. The fossils constitute the most substantial collection of Palaeogene bird bones from Eastern Europe and contribute to a better knowledge of the Paratethyan seabird fauna. Most of the specimens belong to Pelagornithidae (bony‐toothed birds), and two species of very different size can be distinguished. The larger of these is tentatively referred to Dasornis sp., the smaller to Odontopteryx toliapica. The specimens include skeletal elements that were not described for Palaeogene bony‐toothed birds and document previously unknown morphological differences between Palaeogene and Neogene Pelagornithidae. It is argued that the purported crane Eobalearica tugarinovi, from the middle Eocene of Kyrgyzstan, is probably also a bony‐toothed bird. A new genus and species of small Gaviiformes, Colymbiculus udovinchenkoi, is described, which is the earliest fossil record of a loon from Europe, preceding the next oldest specimens by more than 10 myr. The Ukrainian fossils document profound differences between middle Eocene and extant marine avifaunas of Europe, and whereas the middle Eocene Paratethyan avifauna appears to have been similar to that of the North Sea with regard to pelagornithid diversity, the absence of prophaethontids and relative abundance of Gaviiformes may indicate faunistic differences concerning the remaining seabirds.     

New chiropterans from the middle Eocene of Shanghuang (Jiangsu Province,Coastal China): new insight into the dawn horseshoe bats (Rhinolophidae) in Asia

Until recently, the fossil record of Paleogene bats in Asia primarily included extinct families (i.e. ‘Eochiroptera’) from the early Eocene of Vastan in India and from the middle‐late Eocene of the Liguanqiao and Yuanqu basins in central China. Here, we describe a new fauna of Chiroptera from the middle Eocene Shanghuang fissure fillings of China. The fauna includes abundant material referred to a new rhinolophid (Protorhinolophus shanghuangensis gen. and sp. n.), one specimen of a possible rhinopomatid and several indeterminate rhinolophoids. This new bat assemblage constitutes the earliest record of extant families of microbats in Asia. Because it lacks representatives of ‘Eochiroptera’, this Shanghuang bat fauna indicates significant turnover in Asian bat communities. The dental pattern of P. shanghuangensis shows a mosaic of primitive and derived features (‘Eochiroptera’ vs Rhinolophidae dental characteristics), suggesting that this taxon occupies a basal position among the Rhinolophidae. Rhinolophids were already well diversified at the end of the late Eocene in Europe. Interestingly, many dental characteristics of Protorhinolophus are also found in a primitive rhinolophoid taxon, Vaylatsia, from the middle Eocene to late Oligocene of Europe, supporting a close relationship between these taxa. These affinities testify to the widespread Eurasian distribution of rhinolophoids during the Eocene and are consistent with a westward dispersal of the group from eastern Asia to Europe owing to the greater antiquity of Protorhinolophus.     

Evidence for a pre-PETM dispersal of the earliest European crocodyloids

Crocodyloid remains from the late Paleocene of Mont de Berru (France) hosted in the collections of the Muséum National d’Histoire Naturelle (Paris, France) and in the Institut royal des Sciences naturelles de Belgique (Brussels, Belgium) are described for the first time. This material, although fragmentary, can be clearly referred on a morphological basis to Asiatosuchus depressifrons (Blainville, 1855), a species previously reported from several Eocene Belgian localities thanks to abundant material including a nearly complete skeleton. The Paleocene material shares with A. depressifrons the number of alveoli involved in the dentary symphysis, the exclusion of the splenials from the symphysis, and the presence of a distinct depression on the jugal. The fossil remains from Berru represent the oldest European crocodyloid. Along with the alligatoroid Diplocynodon remensis Martin, Smith, de Lapparent de Broin, Escuillié and Delfino, 2014, previously reported from the same locality, the crocodyloid A. depressifrons indicates that these genera reached Europe before the Paleocene–Eocene Thermal Maximum. Although more complete remains from outside Europe are needed to refine phylogenetic hypotheses, according to the currently established fossil record the forerunners of diplocynodontids likely dispersed from North America, whereas those related to Asiatosuchus likely dispersed from Asia.     

Squamate reptiles from the middle Eocene of Lissieu (France). A landmark in the middle Eocene of Europe

In Europe, faunas of squamates (lizards and snakes) from the middle Eocene are very poorly known, with the exception of those from the level MP 16 (latest middle Eocene). From the MP 11-MP 15 interval, squamates were previously reported only from Messel (MP 11, earliest middle Eocene) and from the untere and obere Mittelkohle of Geiseltal (MP 12 and MP 13 respectively) in Germany. The present report describes the middle Eocene assemblage of squamates from Lissieu (France), the first fauna reported from the level MP 14. Whereas fossils from Messel and Geiseltal are mostly articulated skeletons, fossils from Lissieu are represented by disarticulated bones; such fossils may be more easily compared to those from other Cenozoic localities, in which bones are almost always disarticulated. The fauna from Lissieu is more diverse than those from the Geiseltal sites and approximately as diverse as that from Messel as they are presently known; it is comprised of 17 distinct taxa. These taxa cannot be all identified to the species or genus level. They belong to iguanids, gekkonids, lacertids, anguids, thecoglossan platynotans, ophidians incertaesedis, boids, ?tropidophiines, “tropidophiids” incertaesedis, booids incertaesedis, and perhaps russellophiids. The fauna includes several new taxa but only a presumed tropidophiine snake may be named on the basis of the available material. The fauna from Lissieu is a mixture of taxa restricted to the middle Eocene and taxa known from older or younger levels. Taxa shared by Lissieu and the few other localities from the middle Eocene of Europe are rare. This fauna from Lissieu represents a stratigraphical landmark for the middle Eocene.     

New hyaenodonta (Mammalia) from the middle Eocene of Myanmar

The middle Eocene Pondaung Formation in Myanmar has yielded a rich mammalian fauna including several Primate taxa. Hyaenodonta are known by the genera Kyawdawia, Yarshea, Orienspterodon, and two other indeterminate taxa. We describe here new material of Kyawdawia, including some morphological details, a new species of the hypercarnivorous genus Propterodon and an indeterminate species, different from those described earlier in Myanmar, and characterized by a reduction of m3 and would belong to a third lineage with the same evolutionary trend as Galecyon and the Limnocyoninae. The hyainailourines (Orienspterodon) and hyaenodontines (Propterodon) are recorded for the first time in Southeast Asia and these subfamilies appeared in quasi the same time in Europe illustrating probably a profound change in the carnivorous fauna among Laurasia.     

On the early biogeographical history of the African placentals

The Paleocene Adrar Mgorn local fauna recently discovered in the Ouarzazate basin (Morocco) along with several significant Eocene North African faunas, has yielded the oldest known placental mammals of Africa. Contrary to those from the Eocene which are basically endemic, the Adrar Mgorn placentals display affinities with taxa from North‐Tethyan continents and indicate active faunal interchanges between Africa and Europe (and perhaps Asia) during the Cretaceous/Paleogene times. On biogeographical grounds, two dispersal events are suggested as a working hypothesis. The oldest one, exemplified by the presence of paleoryctid and adapisoriculid “insectivores”; in the Moroccan locality, possibly took place by the K/T boundary. The second dispersal event exemplified by the discovery of an omomyid primate and possible hyaenodontid creodonts may have been contemporaneous with the Paleocene/Eocene boundary during which a marine regression is also known.     

First occurrence of Palaeotheriidae (Perissodactyla) from the late–middle Eocene of eastern Thrace (Greece)

A detailed assessment of postcranial fossils collected at Balouk Keui (Thrace, Greece) in the mid-19th Century by the naturalist Auguste Viquesnel enabled us to identify the material as pertaining to Palaeotherium sp., cf. P. magnum, which constitutes the easternmost occurrence of the genus during the Eocene. We have constrained the geographic and stratigraphic provenance of the fossil by reassessing information about Viquesnel's itinerary and observations. Although the exact age of the fossil remains uncertain, the occurrence of a palaeothere in the Thrace Basin during the Eocene indicates a wider geographic distribution for the genus, which had previously been restricted to western and central Europe. The palaeothere of Balouk Keui confirms that the palaeogeographic range of this group included the Balkans during the middle–late Eocene. This discovery also shows that at least intermittent land connections between western Europe, Southeast Asia, and perhaps Central Iran facilitated mammalian dispersal during the middle–late Eocene, before the famous Grande Coupure.     

Amphibians and reptiles from the Early Miocene of the Bardenas Reales of Navarre (Ebro Basin, Iberian Peninsula)

The Lower Miocene deposits of the Bardenas Reales of Navarre (NW Ebro Basin, northern Iberian Peninsula) have yielded a diverse vertebrate fauna, including remains of amphibians and reptiles. These remains occur in several localities in the Tudela Formation. The fossiliferous levels belong to the Biozones MN2b-3 (Biozones Z-A of the Ramblian, i.e., Late Aquitanian to Early Burdigalian in age). The amphibians and reptiles represent at least 13 out of 37 vertebrate species. Amphibians consist of a salamandrid urodele and two or three anurans. All the turtles are cryptodirans and consist of the chelydrid Chelydropsis apellanizi, the testudinids Ptychogaster (Temnoclemmys) bardenensis and Ptychogaster ronheimensis, and a Trionychinae indet. Squamates are represented by the anguid lizard Ophisaurus sp., a non-anguid lacertilian, an amphisbaenian, the erycine boid? Eryx sp., and indeterminate colubrids. Crocodilian remains are assigned to the basal alligatoroid Diplocynodon sp. The fossil associations of the Bardenas Reales of Navarre suggest that the vertebrates lived in the centre of an endoreic basin with stretches of water under intertropical to subtropical climatic conditions.     

Endemism,palaeoendemism and migration: the case for the ‘European endemic’, Mallomonas intermedia

The Synurophyceae is a well-supported clade of ecologically important heterokont algae found largely in freshwater planktonic habitats worldwide, whose members have cell coverings consisting of species-specific siliceous scales overlapped in a highly organized manner. Many synurophytes have been described as endemic and are found only in specific regions of the world. A thriving population of the European endemic, Mallomonas intermedia, was discovered in a remote desert pond situated in the Virgin Valley, Nevada, USA and in a stratigraphic sequence from the middle Eocene fossil locality known as Horsefly in British Columbia, Canada. Both North American finds were closely compared with populations from Europe, confirming the identifications. Before these discoveries, this species was recorded from numerous waterbodies exclusively in Europe, but was lacking from hundreds of sites examined from other continents. Its presence in western North America during the warm middle Eocene confirms that historically this species had a significantly wider distribution and may be best classified as a palaeoendemic. Additional species uncovered from a second fossil locality that are closely related to M. intermedia further support the presence of this lineage in North America during the Eocene. The living population in northern Nevada presents an enigma. Does this remote desert population represent a remnant population that has gone undetected until now, or is it a recent arrival from an unknown region by an unknown vector?     

New chondrichthyan fauna from the Palaeogene deposits of Barmer district,Rajasthan, western India: Age,palaeoenvironment and intercontinental affinities

Field investigations of the Palaeogene sections exposed in Barmer district, Rajasthan (India) led to the discovery of a new fossiliferous horizon in the Padma Rao Open Cast Bentonitic Clay Quarry. The bentonitic clay sequence of this quarry is considered as representing the upper part of the shale, carbonaceous shale, lignite and bentonitic clay succession of the Akli Formation exposed in the Giral Lignite Mine, 2.5 km to its north. Screen-washing of the sediments from a fossiliferous level within the bentonitic clay of the new site has yielded nine taxa of sharks and batoids: Squatiscyllium nigeriensis White, 1934, Ginglymostoma sokotoense White, 1934, Ginglymostomatidae gen. et sp. indet., Brachycarcharias sp., Abdounia recticona Winkler, 1873, Premontreia (Oxyscyllium) peypouqueti Noubhani et Cappetta, 1997, Eotorpedo hilgendorfi Jaekel, 1904, Coupatezia sp. cf. C. danica, and Myliobatidae gen. et sp. indet. The fauna recovered is mainly represented by nearshore shallow marine forms. This fauna is quite different from that of Palaeocene Fatehgarh Formation, the Lower Eocene Kapurdi Formation (Barmer Basin), the Khuiala Formation (Jaisalmer Basin) and the Cambay Shale (Cambay Basin), and favours a late Palaeocene (Thanetian) age for the upper part of the Akli Formation. A nearshore shallow marine palaeoenvironment is favored for the investigated stratigraphic horizon on the basis of the selachian fauna. Majority of the fossil shark species described from the Padma Rao quarry section are known from a number of sites in Africa, western Europe, Asia and southeastern USA, suggesting that free faunal interchanges were possible between the western Tethys and Indo-Madagascan faunal provinces during the late Palaeocene.     

The “miacids” (Carnivoraformes,Mammalia) from the Early Eocene locality of Le Quesnoy (MP7, France); first occurrence of Vassacyon in Europe

We describe here “miacid” taxa from the Early Eocene Paris Basin locality of Le Quesnoy (Oise, France). We describe the new species Vassacyon taxidiotis, the first European record of this genus. The other “miacids” identified from Le Quesnoy are Miacis latouri and Gracilocyon solei. The P4 of G. solei is described here for the first time. Its morphology (e.g., wide protocone, short postmetacrista) supports a close relationship with Miacis rundlei from Abbey Wood (MP8 + 9, England). The latter species is therefore classified as Gracilocyon rundlei. Three new tooth positions are known for Miacis latouri: P4, p4 and m2. They support its reference to Miacis. These specimens imply that the European species is more basal than the North American species. The fauna from Le Quesnoy shares with Dormaal the presence of Miacis latouri and Gracilocyon solei, but the “miacid” fauna from Le Quesnoy also contains Vassacyon taxidiotis. The presence in Le Quesnoy of the two former taxa supports a reference to MP7 level of the French locality. The presence of three distinct genera in European localities show that the “Miacidae” were diversified in Europe, as previously observed in North America. The genera Gracilocyon, Miacis, and Vassacyon probably dispersed from Europe to North America during the Palaeocene/Eocene boundary.     

Vertebrates from the Pontian of the Shkodova Gora locality (Northwestern Black Sea Region,Upper Miocene)

Remains of fishes, turtles, birds, and mammals are described from the Upper Miocene (MN13) Shkodova Gora locality (Ukraine, Odessa Region), the only representative locality of Pontian large vertebrates in the northwestern Black Sea Region. Asiatic fish and avian taxa are recorded for the first time in the Upper Miocene of Europe. The Shkodova Gora ichthyofauna includes eight freshwater fish species (belonging to Acipenseridae, Cyprinidae, Siluridae, and Percidae). Taxa that have previously been recorded only in Western Siberia (Abramis bliccoides, Perca lepidopoma), eastern Kazakhstan, and the Altai Mountain (Rutilus tungurukensis) are revealed. Acipenser gueldenstaedtii is recorded for the first time in the Upper Miocene of Eastern Europe. The occurrence of the genus Ctenopharyngodon, a member of the Chinese plain faunal assemblage is of great interest. The Shkodova Gora avifauna (seven taxa) is mostly composed of waterfowl, including Phalacrocorax mongoliensis, which has previously been known only from the Lower Pliocene of Mongolia. The finding of Proanser major is the youngest record of this species in the fossil record. Mammals of the Shkodova Gora association include a hare resembling Trischizolagus dumitrescuae, Palaeoryx cf. pallasi, and Hipparion cf. moldavicum, which have not been mentioned among Pontian taxa of the northern Black Sea Region. Pontian paleolandscapes in the area of the Shkodova Gora locality are reconstructed.     

Fossil crocodylid remains from Scontrone (Tortonian,Southern Italy) and the late Neogene Mediterranean biogeography of crocodylians

New fossil remains from the Tortonian of Scontrone (Abruzzo, Southern Italy) provide further information on crocodylids with a generalized rostral morphology that inhabited the Mediterranean area during the late Miocene. Particularly informative is the nearly complete right dentary SCT 276. Thanks to the fact that the third and fourth alveoli are clearly separated by a bony septum and are markedly different in size, the fourth being much larger than the third, it is possible to exclude that SCT 276 belonged to the alligatoroid Diplocynodon and to tentatively refer it to cf. Crocodylus sp. This genus has been previously identified on a phylogenetic basis in the same Apulo-Abruzzi palaeobioprovince (Gargano Terre Rosse, Messinian-Zanclean, Crocodylus sp.), and it is likely present also in the Tusco-Sardinian palaeobioprovince (Monte Bamboli, Tortonian, cf. Crocodylus sp.). SCT 276 currently represents the oldest possible evidence of the presence of Crocodylus; it proves that this taxon could have already reached Europe during the Tortonian, well before the Messinian Salinity Crisis that is traditionally considered as the event that caused several trans-Mediterranean dispersals. Furthermore, it is tempting to associate the absence of alligatoroids and the presence of crocodylids in these palaeobioprovinces (actually systems of islands) to the different behavioural, morphological and physiological traits of extant alligatorids and crocodylids, which render rather salt-intolerant the former and salt-tolerant the latter.     

The parrots (Aves: Psittaciformes) from the Middle Miocene of Sansan (Gers,Southern France)

The occurrence of Archaeopsittacus sp. (Psittaciformes) in the fossil deposits of Sansan (France) is reported, testifying to a survival of this genus in the Middle Miocene, as Archaeopsittacus verreauxi was described from the Early Miocene of Saint-Gérand-le-Puy (France) and was recorded only from its type locality. The data discussed here indicate the presence of two parrot species from Sansan, as it is the type locality of another parrot species, Pararallus dispar, only known from this locality. The differences between the humeri of these two taxa are described in detail, together with the differences from the other European fossil parrot species. The presence of more than one species of parrot in the same locality is not rare, but in Europe it is recorded in Sansan for the second time. Evidence for parrots also confirms the paleoenvironmental reconstruction of Sansan, as parrots are primarily arboreal species. Archaeopsittacus also represents one of the few common elements between the Early and Middle Miocene European bird assemblages.