An Eocene tomistomine from peninsular Thailand

Skull and mandibular elements of a tomistomine crocodilian are described from the late Eocene to early Oligocene lignite seams of Krabi, peninsular Thailand. The Thai tomistomine is a longirostrine form characterized by a rostrum/skull ratio of about 0.6; a mandibular symphysis reaching the level of the eleventh alveolus; a deep participation of the splenial in the symphysis to the level of the ninth alveolus; an enlarged fifth maxillary alveolus; long nasals reaching the premaxillae at the level of the fifth maxillary alveolus but not contributing to the external nares; undivided posterior processes of premaxillae; a short prefrontal, excluded by a longer lacrimal from the posteromedial margin of maxillae; vomers visible on the palate. A phylogenetic analysis recovers the Thai specimen among derived tomistomines, on the stem lineage of the extant Tomistoma schlegelii, as a close ally of Maomingosuchus petrolica, a late Eocene tomistomine from southern China. The present recognition of Maomingosuchus sp. in late Eocene Thai deposits expands the distribution of Eocene tomistomines from southern China to the tropics. The origin of Tomistominae in Asia is briefly discussed.     

The oldest representative of Entelodontoidea (Artiodactyla,Suiformes) from the Middle Eocene of Khaichin Ula II,Mongolia, and some evolutionary features of this superfamily

The oldest representative of the family Entelodontidae (suborder Suiformes), Proentelodon minutus gen. et sp. nov, is described from the Middle Eocene Khaichin Ula II Fauna in Mongolia. In addition to the genus Proentelodon, the new subfamily Proentelodontinae includes the genus Eoentelodon from the Middle-Upper Eocene of China, Mongolia, and Kazakhstan. The new data bring clues to the origin and systematic position of Entelodontidae, features of their evolution, dispersal, and changes in taxonomic diversity. They also elucidate the early evolutionary stages of Suiformes.     

First fossil record of the genus Nageia (Podocarpaceae) in south China and its phytogeographic implications

The first fossil record of the genus Nageia Gaertner from the Tertiary of southern China is reported. Nageia hainanensis sp. nov. is described from the Eocene Changchang Formation of Hainan Island. The extant species of Nageia are widely distributed in southeastern Asia, from northeastern India to southeastern China and southern Japan, and southward through Indochina to the Malay Archipelago and New Guinea. The fossil evidence of Nageia has been discovered from the Lower Cretaceous of the Far East of Russia and Japan. The distribution of both modern and fossil species of Nageia indicates that this genus could have originated in the northeastern part of Asia in the Early Cretaceous and spread to south China at least in the Eocene. Discovery of Nageia fossil record in south China provides important evidence for the geological and phytogeographic history of the genus.     

New adapiform primate of Old World affinities from the Devil's Graveyard Formation of Texas

Most adapiform primates from North America are members of an endemic radiation of notharctines. North American notharctines flourished during the Early and early Middle Eocene, with only two genera persisting into the late Middle Eocene. Here we describe a new genus of adapiform primate from the Devil’s Graveyard Formation of Texas. Mescalerolemur horneri, gen. et sp. nov., is known only from the late Middle Eocene (Uintan) Purple Bench locality. Phylogenetic analyses reveal that Mescalerolemur is more closely related to Eurasian and African adapiforms than to North American notharctines. In this respect, M. horneri is similar to its sister taxon Mahgarita stevensi from the late Duchesnean of the Devil’s Graveyard Formation. The presence of both genera in the Big Bend region of Texas after notharctines had become locally extinct provides further evidence of faunal interchange between North America and East Asia during the middle Eocene. The fact that Mescalerolemur and Mahgarita are both unknown outside of Texas also supports prior hypotheses that low-latitude faunal assemblages in North America demonstrate increased endemism by the late middle Eocene.     

Bombacaceae pollen from the Indian Tertiary sediments and its bearing on evolution and migration

Several pollen species recovered from the Indian Tertiary sediments have close similarities with pollen of extant Bombacaceae. These morpho species are included in the morphogenera Lakiapollis Venkatachala and Kar, Dermatobrevicolporites Kar, Bombacacidites Couper, Tricolporocolumellites Kar and Retitribrevicolporites Kar. Nine species are re-described on the basis of original type material or other specimens. The fossil pollen belongs to the Bombax- and Durio-types. The small-sized Bombax-type pollen (e.g., Eriotheca pollen) has been documented in India from the early Palaeocene to the Miocene while large-sized pollen similar to that of Bombax appeared in the Oligocene and continued up to the Present. The lobed variety of the Bombax-type is rare and occurs only in the Pleistocene. Palynological evidence suggests a migration of Bombacaceae producing Bombax-type pollen to India from Africa. Durio-type pollen appeared in the late Palaeocene and continued to dominate until the early Eocene. In southern India, this pollen remained frequent during Oligocene/Miocene times, whereas in other areas, it became rare at the end of the Eocene. The earliest (late Palaeocene) record suggests an origination of Bombacaceae producing Durio-type pollen in India and a subsequent migration to Southeast Asia.     

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.     

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.     

Simimys and origin of the cricetidae (Rodentia: Muroidea)

Simimys is a late Eocene and earliest Oligocene genus that shares dipodoid (zygoma) and muroid (dental) characters. The Oligocene record of dipodoid rodents includes Plesiosminthus from middle Oligocene deposits of Asia and late Oligocene deposits of Europe. The Oligocene record of muroid rodents includes at least two genera (Eucricetodon and (Cricetops)) from Asia, six genera (Eucricetodon, Pseudocricetodon, Melissiodon, Paracricetodon, Heterocricetodon and Adelomyorion)) from Europe, and three genera (Eumys, Scottimus and Nonomys)) from North America. The known record, as given above, suggests that Siminys is the earliest and most primitive genus with muroid affinities; it also implies that muroid rodents were derived from unknown Eocene dipodoid rodents.The Oligocene cricetid rodents display progressive expansion and inclination of the anterior plate of the zygoma. These changes in the zygoma probably reflect evolutionary stages in the development of a myomorphous zygoma from an hystricomorphous zygoma. Changes in the zygoma apparently took place at different rates and times in Asia. Europe, and North America; they probably reflect differenciation of Cricetodontinae in Palaearctica, and Eumyinae in Nearctica during the Oligocene.     

First partial cranium of Togocetus from Kpogamé (Togo) and the protocetid diversity in the Togolese phosphate basin

Earliest cetaceans (whales) originated from the early Eocene of Indo-Pakistan, but the group dispersed through most of the oceans of the planet by the late middle to late Eocene. This late Eocene global distribution indicates that important dispersal events took place during the middle Eocene (Lutetian), a globally undersampled time interval that is well documented in the Togolese phosphate series. We report here the first discovery of a partial cetacean cranium from middle Eocene deposits of Togo (West Africa). A 3D model of the cranium and teeth was reconstructed in order to reveal hidden anatomical features. The dental and cranial characteristics of the Togolese specimen recall those of protocetid taxa described in Africa, Asia, and North America, but also display significant differences. In particular, we show that the new specimen shares a number of morphological features with the Togolese taxon Togocetus. Such a hypothesis is further supported by a cladistic analysis including 45 taxa and 167 morphological characters, which recovers the new specimen close to Togocetus as the first offshoot of protocetids. Phylogenetic analysis including all the protocetids remains of Kpogamé confirms the singular diversity of the Togolese phosphate basin, and enables to examine potential connections with faunas from contemporaneous localities in Africa.     

Zhailimeryx, A New Lophiomerycid Artiodactyl (Mammalia) from the Late Middle Eocene of Central China and the Early Evolution of Ruminants

A new genus and species of primitive selenodont artiodactyl, Zhailimeryx jingweni, occurs in the late middle Eocene Zhaili Member of the Heti Formation, Yuanqu Basin, Shanxi Province, China. A phylogenetic analysis of dental characters suggests that Zhailimeryx is closely related to Lophiomeryx and other lophiomerycid ruminants of Asia and Europe. Zhailimeryx and other Eocene records of Lophiomerycidae from Asia support an Asian origin for this family, followed by dispersal into Europe both before and after the Grande Coupure. Morphological evidence from Zhailimeryx suggests that ruminant artiodactyls had already undergone substantial diversification prior to the late middle Eocene, and it reopens the issue of whether the higher taxa Tragulina and Pecora are valid expressions of ruminant phylogeny.     

A new basal tapiromorph (Perissodactyla, Mammalia) from the middle Eocene of Myanmar

A new genus and species of tapiromorph, Skopaiolophus burmese nov. gen., nov. sp., is described from the middle Eocene Pondaung Formation in central Myanmar. This small form displays a striking selenolophodont morphology associated with a mixture of primitive “condylarthran” dental characters and derived tapiromorph features. Skopaiolophus is here tentatively referred to a group of Asian tapiromorphs unknown so far. The occurrence of such a form in Pondaung suggests that primitive tapiromorphs might have persisted in southeast Asia until the late middle Eocene while they became extinct elsewhere in both Eurasia and North America.     

First record of non-marine ostracods from the Paleogene “hamadian deposits” of Méridja area,west of Bechar (southwestern Algeria)

A new non-marine ostracod fauna from the Paleogene “hamadian deposits” outcropping west of Bechar (southwestern Algeria) has been recovered from lacustrine to fluvial deposits of the Oued Méridja section and fluvial deposits on the southern edge of the Hamada de Méridja section. Recently, these sections have been dated as late Thanetian – early Ypresian (latest Paleocene to earliest Eocene) and Ypresian – earliest Lutetian (early to earliest middle Eocene), respectively, based on charophytes. The associated ostracod fauna recovered consists of relatively mostly moderately to badly preserved specimens and comprises 14 taxa, none of which could be identified to species level in view of its poor state of preservation; we have nevertheless been able to identify and describe the following taxa: Herpetocypris sp., Cyprinotus? sp., Heterocypris? sp. 1 and sp. 2, Cypris? sp., Ilyocypris sp., Cytheroidea indet. sp. 1 and sp. 2, Limnocytheridae indet. sp. 1, Cypridoidea indet. sp. 1, Cyprididae indet. sp. 1, and Ostracoda indet. sp. 1, 2 and 3. Only Heterocypris sp. 1 occurs in both sections. Although the fauna can as yet not be related to the few other contemporaneous faunas reported from the wider palaeogeographic area, it adds important new information to our poor knowledge on Eocene non-marine ostracods in North Africa and southern Europe. The Méridja sections and area are promising regarding the discovery of more, better preserved material and further studies, and one main limitation to the correlation of the fauna is the hitherto insufficient taxonomic knowledge on many faunal elements of Eocene non-marine ostracods to which our section contributes considerably.     

Oldest known Varanus (Squamata: Varanidae) from the Upper Eocene and Lower Oligocene of Egypt: support for an African origin of the genus

Abstract: A large collection of lizard vertebrae from northern Africa represents the oldest unambiguous occurrence of the genus Varanus. The fossils come from late Eocene and early Oligocene freshwater deposits of the Fayum, Egypt, an area noted for many significant primate finds. The recovery and identification of this material indicate that the genus Varanus arose in Africa, before dispersing to Australia and Asia. This dispersal occurred prior to the early to mid‐Miocene, by which time fossil Varanus are known from Australia and Eurasia. Although the dispersal route remains unknown, the lizard material reported here supports the hypothesis that a corridor existed allowing freshwater and terrestrial organisms to cross from Africa to Asia.     

The taxonomy,chronostratigraphy and paleobiogeography of glyptosaurine lizards (Glyptosaurinae,Anguidae)

Glyptosaurine lizards (Glyptosaurinae, Anguidae) are an extinct group of heavily armored lizards known from North America, Europe and Asia. Glyptosaurine lizards, taxa that possess fully developed tuberculated dermal armor, appear to have been established in North America by late early Puercan time (To3). “Proxestops,” a taxon distinguished by a combination of vermiculate and tuberculated osteoderm sculpturing, is considered to be a non-glyptosaurine, a sister taxon of the Glyptosaurinae. Known from only fragmentary remains, its wide chronostratigraphic distribution suggests that “Proxestops” is a form genus that, in all probability, represents more than one taxon, that ranges from the middle Paleocene to the early Eocene of North America. Moreover, the taxa Odaxosaurus piger, Parodaxosaurus sanjuanensis and “Proxestops” are best considered “proto-glyptosaurines”. “Melanosaurins” and glyptosaurins were well-established by the early Eocene, especially in North America, and are here documented by their type species and chronostratigraphic levels. Both tribes are present in Europe (MP7), too, but the record is not as estensive as that of North America. The North American taxon Gaultia silvaticus (Wa0) is transitional between a “melanosaurin” and glyptosaurin. Because it lacks the well-defined hexagonal osteoderms that characterize the Glyptosaurini, it is removed from that group and considered to be a “melanosaurin”. The “melanosaurin” taxon “Xestops” savagei (Wa4–Wa6) cannot be referred to Xestops (Br2) based on non-corresponding elements and because superficial similarity does not justify assignment to this taxon. Arpadosaurus sepulchralis (Wa6?), whose holotype is a fragmentary right frontal, is considered a subjective junior synonym of A. gazinorum, based on minor differences in the epidermal scale pattern that probably represent individual variation. “Glyptosaurus” agmodon (Wa6?), based on a partial right maxilla, cannot be referred to Glyptosaurus (sensu stricto), and the material upon which this taxon is based bears strong resemblance to material identified as cf. “?Paraglyptosaurus” yatkolai (Wa5–Wa6). “Glyptosaurus” rhodinos (Wa5) is based on an incomplete parietal, and its reference to Glyptosaurus is considered problematic. Eoglyptosaurus donohoei (Wa7) is probably valid and is re-established here. Glyptosaurus (sensu stricto) is known solely from the middle Eocene (Br2) by G. sylvestris. Dimetoposaurus wyomingensis (Br3) is removed from Xestops vagans because its synonymy was based on superficial similarities. Helodermoides tuberculatus, the largest and last glyptosaurin (Ch3), is restricted to the Chadronian of North America. Only the “melanosaurin” Peltosaurus granulosus (Or2–Or3), which includes the species P. abbotti, seems to have crossed the Eocene-Oligocene boundary, and appears to be largely restricted to the Orellan, but extended into the Arikareean. European glyptosaurines are also represented by both glyptosaurins and “melanosaurins” early in the Eocene (MP7). Placosauriops-like “melanosaurins” are known from Dormaal (MP7), and the glyptosaurin taxon?Placosaurus ragei occurs at the same level. “Placosauriops abderhaldeni” has been identified from the Grube Messel (MP11), but this assignment remains dubious because the species has not been adequately diagnosed, and the holotype species is from the Geiseltal (MP13), which is some 4.5 million years younger. Placosauriops weigelti (MP13) is the only valid species of this genus. Paraxestops stehlini (MP14) is not referable to the North American taxon Xestops, and its relationship to Placosauriops has not been studied. The late Eocene glyptosaurins Placosaurus estesi (MP17) and P. rugosus (MP18) are the last glyptosaurines known from Europe and appear to have gone extinct at the Eocene-Oligocene boundary, casulties perhaps of the “Grande Coupure”. Asian glyptosaurines are known solely from one species, Stenoplacosaurus mongoliensis, from the middle Eocene (Sharamurunian) of China. Glyptosaurines most likely originated in North America, diversified by late Paleocene time, and rapidly spread across the North Atlantic into Europe by the early Eocene. Both “melanosaurins” and glyptosaurins took a foothold in Europe by the early Neustrian, but the glyptosaurins, aside from one occurrence (Dormaal, MP7), were conspicuously absent for most of Neustrian through early Robiacian time. In North America, glyptosaurins diversified during the early and middle Eocene, while in Europe small “melanosaurins” were a prominent part of the paleoherpetofauna, and glyptosaurins are unknown for most of the Neustrian through the Geiseltalian, in both the fossilferous Lagerstätten of Messel and Geiseltal. Stenoplacosaurus is the only known glyptosaurin glyptosaurine from Asia, and its abrupt appearance during the late Eocene suggests the possiblity of a Beringian dispersal from North America into Asia.     

Crocodylian assemblage from the middle Eocene Ikovo locality (Lugansk Province,Ukraine), with a discussion of the fossil record and geographic origins of crocodyliform fauna in the Paleogene of Europe

The known fossil record of crocodyliforms in Europe during the Paleogene is significantly biased, in that the fauna of Western Europe is far better sampled and understood compared to that of Eastern Europe. We describe in detail all known crocodyliform remains from the middle Eocene (Lutetian) Ikovo locality in Ukraine. We conclude that at least two taxa were present: a moderate to large-sized Tomistominae indet. similar to the basalmost known tomistomines, and the small-sized basal alligatoroid cf. Diplocynodon sp. Despite its scarcity, this is the first basal alligatoroid material reported from Eastern Europe (as part of post-Soviet countries) and the easternmost record of diplocynodontines in Europe so far. An allegedly freshwater cf. Diplocynodon sp. contributes a rare faunal element to the vertebrate assemblage of the Ikovo locality, otherwise dominated by resident or facultative marine taxa. The fossil record and historical paleobiogeography of crocodyliforms from the Paleocene and Eocene of Europe are reviewed. As it has been already known, the middle Eocene fauna of crocodyliforms proves to be taxonomically diverse and complex. Its constituent lineages geographically originated in Asia or North America (Diplocynodontinae, Asiatosuchus-like crocodyloids, Planocraniidae), North America (derived alligatorines), Africa (Tomistominae), and Gondwana (ziphodont mesoeucrocodylians Iberosuchus and Bergisuchus), with possible subsequent speciation in Europe. We propose a novel hypothesis of Asian origins of European diplocynodontines, which will be explicitly tested in future studies. The revealed similarities between crocodylians and turtles from the Ikovo locality and those from Western Europe support the presence of a single Pan-European biogeographical zone during the middle Eocene, distinct from that of Asia.     

A goose-sized anseriform bird from the late Oligocene of France: the youngest record and largest species of Romainvilliinae

We describe a new anseriform bird from the late Oligocene of Saint-André, Marseille, in southern France. Saintandrea chenoides, gen. et sp. nov. is the first avian species reported from the locality, which is well known for its mammalian fossils. The new species belongs to the extinct Romainvilliinae and represents the latest occurrence of the taxon, which was before only known from the late Eocene and early Oligocene of Europe. S. chenoides is also the largest species of Romainvilliinae and increases the known morphological diversity of the taxon. The identification of a goose-sized representative of the Romainvilliinae in the late Oligocene of Europe raises the possibility that some of the large late Paleogene or early Neogene Anseriformes with uncertain phylogenetic affinities also belong to this taxon.     

New macrostomatan snake from the Paleogene of northwestern Argentina

The lower Eocene Lumbrera Formation in Salta province, northwestern Argentina, outstands for providing snake remains from a non-Patagonian Paleogene site. The material consists of articulated precloacal vertebrae that represent a new medium-sized macrostomatan snake, namely Amaru scagliai nov. gen., nov. sp. The vertebral characters of Amaru scagliai nov. gen., nov. sp., suggest affinities with advanced clades, which is consistent with the recognition of derived macrostomatans in the early Paleocene of Bolivia and early Eocene of Brazil. The new snake confirms the presence of macrostomatan snakes in South America as early as the Eocene and suggests that the southern continents may have played an unsuspected role in the origin and evolution of advanced macrostomatans during the earliest Cenozoic.     

A Review of the Fossil Record of the Genus Itea (Iteaceae, Saxifragales) with Comments on its Historical Biogeography

Itea is a genus of about 20 species of trees and shrubs that are today native to southeastern North America, eastern Asia, and eastern Africa. In this paper, I review the fossil record of Itea, which is based on four types of fossils: diporate, psilate pollen attributed to Itea or the dispersed pollen genus Iteapollis; carpofossils representing fruits and seeds attributed to Itea europaea; flowers preserved in amber and assigned to Adenanthemum iteoides; and leaf impressions attributed to Itea. The distributions of these fossils indicate that Itea was present in western North America from the early Eocene to Miocene, in eastern North America beginning no later than the early Miocene, and in western Eurasia from the late Eocene to Pliocene. Only one datapoint is known from eastern Asia; it is early Miocene in age. Based on the fossil record, it can be inferred that Itea crossed between continents over both the Bering Land Bridge and North American Land Bridge, and that it reached Africa from Europe via Anatolia. Thus, it is predicted that the sole extant North American species, I. virginica, may be most closely related to the sole extant African species, I. rhamnoides. The potential application of Itea fossils to calibrating phylogenetic trees generated from molecular sequence data is also discussed.     

Winged Fruits of Deviacer in the Oligocene from the Ningming Basin in Guangxi,South China

Deviacer guangxiensis Chen & Manchester sp. nov. is described based on asymmetric samaras from the Oligocene Ningming Formation in Guangxi, South China, representing the first documentation of Deviacer fossils in Asia. The Oligocene species, with relatively large fruits, represents the youngest record of the genus so far known; all other records are from the Paleocene and Eocene, or late Eocene—early Oligocene in western North America and Europe. It indicates that the extinct genus, Deviacer, was widely distributed in the northern hemisphere during the Paleogene.