Additional vertebral material of Thaumastophis (Serpentes: Caenophidia) from the early Eocene of India provides new insights on the early diversification of colubroidean snakes

The Ypresian Cambay Shale Formation at Vastan, Mangrol, and Tadkeshwar lignite mines in Gujarat, western India, has yielded a rich vertebrate fauna including madtsoiid, palaeophiid, booid, and colubroidean-like snakes. The latter are particularly abundant, but their systematic affinities are difficult to resolve. Here we describe new specimens of the colubroidean-like snake Thaumastophis missiaeni, including anterior, middle, and posterior trunk vertebrae, as well as caudal vertebrae. The combination of primitive and derived caenophidian and colubroidean vertebral characters confirms Thaumastophis as the earliest known stem-colubriform snake while Procerophis, from the same beds, is more derived and considered to represent a crown-Colubriformes. Additionally, Thaumastophis shares with Renenutet enmerwer from the late Eocene of Egypt a unique combination of vertebral characters that suggests an exchange with North Africa was possible along the southern margin of the Neotethys. We erect the new family Thaumastophiidae for Thaumastophis and Renenutet on the basis of their shared derived vertebral morphology.     

Middle Jurassic Porifera from Kachchh, western India

From the Late Bathonian sponge biofacies at Jumara Dome, Kachchh (western India) 11 species of ‘lithistid’, hexactinellinid and calcarean sponges are described. New taxa are the order Sigmatospirida, the genusJumarella, and the speciesJumarella astrorhiza, Mastosia rhytidodes, Radicispongia kraspedophora, andHexactinella prisca. The diverse sponge assemblage is associated with a rich fauna of epibenthic bivalves and brachiopods and formed meadows on fine-grained carbonate substrates. The sponge meadows grew on a carbonate ramp at the lower end of the photic zone, in quiet waters below storm wave base. The rate of sedimentation exceeded that of sponge production. This prevented the development of reef-like bodies. In contrast to Mesozoic sponge reefs, growth of the sponge meadows appears to have been confined to the regressive phases of small transgressive-regressive cycles.     

The oldest Jurassic cyathophorid coral (Scleractinia) from siliciclastic environments of the Kachchh Basin, western India

Cyathophora Michelin, 1843, hitherto well known from the Upper Jurassic and Cretaceous, has been found in the Middle Jurassic (Bajocian) of the Kachchh Basin, western India. Eleven specimens ofCyathophora bourgueti (Defrance, 1826) from the Babia Cliff Sandstone member of the Kaladongar Formation, exposed along the northern scarp of the Kala Dongar, Pachchham Island, Kachchh, are described and illustrated as the earliest Jurassic record of the family CyathophoridaeVaughan & Wells, 1943. It is suggested that the monospecific occurrence ofCyathophora bourgueti was controlled by salinity.      

Neocrassinid bivalves (Heterodonta) from the Middle Bathonian (Jurassic) of Kachchh,Western India

Six bivalve taxa belonging to the genusNeocrassina Fischer have been described from Middle Bathonian (Middle Jurassic) rocks of Kachchh, Western India. Of these,Neocrassina (N.) ovata (Smith), TV.(N.) robusta (Etallon) andN. (N.) paphia (d?Orbigny) have been recorded previously from various parts of Europe, whileNeocrassina (N.) pandeyi Maithani is an endemic faunal element as areNeocrassina (N.) sp. A andNeocrassina (N.) sp. B. The generic and sub-generic characters ofNeocrassina are reviewed as are affinities of the taxa with closely related forms from India and other parts of the world.     

Fossil ectomycorrhizae from the Middle Eocene

Fossil ectomycorrhizae were found recently among permineralized plant remains in the middle Eocene Princeton chert of British Columbia. The ectomycorrhizae are associated with roots of Pinus and have a Hartig net that extends to the endodermis, a pseudoparenchymatous mantle, and contiguous extramatrical hyphae that are simple-septate. The mycorrhizal rootlets lack root hairs and dichotomize repeatedly to form large, coralloid clusters. Reproductive structures are absent. Based on the morphological characteristics, and the identity of the host, the closely related basidiomycete genera Rhizopogon and Suillus are suggested as comparable extant mycorrhizal fungi. These exquisitely preserved specimens represent the first unequivocal occurrence of fossil ectomycorrhizae and demonstrate that such associations were well-established at least 50 million years ago.     

Reassessment of the Early–Middle Eocene biomagnetochronology based on evidence from the Gorrondatxe section (Basque Country, western Pyrenees)

Thirteen Lower–Middle Eocene (Ypresian–Lutetian) successions, including the Gorrondatxe section in the western Pyrenees, show biomagnetostratigraphic correlation schemes that do not agree with the current standard framework. The main discrepancy concerns the position of the boundary between planktonic foraminiferal Zones P9 (=E7, approximately) and P10 (=E8, approximately), which was thought to occur within calcareous nannofossil Subzone CP12a and at the boundary between magnetic polarity Chrons C22n and C21r. However, in the differing correlation scheme the boundary between Zones P9 (=E7) and P10 (=E8) occurs close to the base of Subzone CP13a and to the boundary between Chrons C21n and C20r. An attempt at a new Ypresian–Lutetian boundary biomagnetochronology is made based on data from the Gorrondatxe section, which shows that the boundary between Zones P9 (=E7) and P10 (=E8) is 3.1 Myr younger than hitherto considered. Therefore, the duration of the Early Eocene, most commonly defined according to this planktonic foraminiferal zonal boundary, has generally been underestimated over the last four decades.     

Systematics and diversity of the giant soft-shelled turtles (Cryptodira,Trionychidae) from the earliest Eocene of Belgium

In 1909, the famous paleontologist Louis Dollo identified two putative new species of giant soft-shelled turtles from the lowest Eocene record of Belgium, ‘Trionyx erquelinnensis’ and ‘Trionyx levalensis’, from Erquelinnes and Leval, respectively. However, these proposals did not meet the requirements of the International Code of Zoological Nomenclature, so they were considered as nomina nuda. The information on these specimens or about any other specimen of this lineage of giant turtles from the Belgian record is currently extremely limited. Relatively scarce material from giant trionychids has been described for the lower Eocene record of other European regions. Considering the available information, all the European material has recently been recognized as belonging to the genus Axestemys, which has a North American origin, and possibly attributable to a single species, Axestemys vittata, which currently lacks a diagnosis. Numerous and well-preserved Belgian specimens are deposited in the Royal Belgian Institute of Natural Sciences. In addition to the cited individuals from Erquelinnes and Leval, additional specimens from both localities, as well as others from Orp-le-Grand, are part of this collection. These specimens, found between the decades of 1910 and 1930, have been recently restored, and their study is carried out here. The presence of Axestemys vittata in Belgium (in Leval and Orp-le-Grand) is confirmed. Knowledge about this species is significantly improved, and a diagnosis is proposed. However, the hypothesis proposed by Dollo is here confirmed, this species being not the only identified in the Belgian record. So, Axestemys erquelinnensis nov. sp. is defined based on the carapace from Erquelinnes known by Dollo, suggesting that the genus probably reached Europe during the Paleocene Eocene Thermal Maximum.     

New material of Diacodexis (Mammalia,Artiodactyla) from the early Eocene of Southern Europe

Diacodexeidae are the first representatives of Artiodactyla in the fossil record. Their first occurrence is at the very base of the Ypresian (earliest Eocene, 56.0 Ma) with Diacodexis, a genus well diversified during the early Eocene in Europe, especially during the MP7–MP8 + 9 interval. However, most of European species are documented by scarce material, retrieved from single localities. In this work, we describe new Diacodexis material from ～MP7 and ～MP8 + 9 localities of Southern Europe, including material of D. antunesi from Silveirinha, considered as the most primitive European Diacodexis species, and material from three localities from Southern France (Fordones, Palette, and La Borie). The new material documents Diacodexis premolar morphology and deciduous dentition which bear potentially important phylogenetic information, as well as astragali, including a specimen from Silveirinha that constitutes the earliest occurrence of an astragalus of the genus Diacodexis in the European fossil record. Investigation of the enamel microstructure reveals that early European species had a simple enamel pattern with one-layered Schmelzmuster composed of ‘basic’ radial enamel only, instead of the two-layered Schmelzmuster (thin radial enamel + thick layer of Hunter-Schreger bands) observed on North American species and so far considered to represent the primitive condition within Artiodactyla. In accordance with previous studies, our observations highlight that Diacodexis gigasei from Belgium is morphologically closer to the North American species D. ilicis than to D. antunesi from Portugal. The latter species, together with D. aff. antunesi from Fordones, appears to be morphologically closer to the Asiatic taxa D. indicus and D. pakistanensis. Finally, we found numerous similarities between D. cf. gigasei from Palette and D. gigasei, a result that challenges the intra-European provincialism that characterizes the earliest Ypresian. Diacodexis gigasei could be one of the rare species shared by the northwestern and southwestern European bioprovinces.     

A new species of Dyrosaurus (Crocodylomorpha, Dyrosauridae) from the early Eocene of Morocco: phylogenetic implications

A new dyrosaurid is described from the Ypresian of the phosphatic deposits of the Oulad Abdoun Basin of Morocco. It is based on numerous cranial and postcranial remains, allowing an almost complete reconstruction. This new Dyrosaurus species, Dyrosaurus maghribensis sp. nov. , is currently only known from Morocco. It differs from D. phosphaticus , present in contemporaneous levels of Algeria and Tunisia, by several autapomorpies, including a smooth dorsal margin of the parietal and widely opened choanae. A phylogenetic analysis, using 47 taxa and 234 morphological characters, shows the dyrosaurids as the sister taxon of pholidosaurids, which include Elosuchus , Sarcosuchus , Terminonaris and Pholidosaurus , and the thalattosuchians. Goniopholididae is a non-monophyletic group; however, if dyrosaurids are not included in the analysis, the result differs and the goniopholidids form a distinct clade. If Thalattosuchia is excluded, both Goniopholididae and Pholidosauridae become paraphyletic assemblages. Thus, phylogenetic problems remain with respect to longirostrine clade, and more attention should be paid to resolving their evolutionary relationships amongst the crocodyliforms.  © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society , 2006, 148 , 603–656.     

The Vermetidae of the Gulf of Kachchh,western coast of India (Mollusca,Gastropoda)

Coral reefs are often termed underwater wonderlands due to the presence of an incredible biodiversity including numerous invertebrates and vertebrates. Among the dense population of benthic and bottom-dwelling inhabitants of the reef, many significant species remain hidden or neglected by researchers. One such example is the vermetids, a unique group of marine gastropods. The present study attempts for the first time to assess the density and identify preferred reef substrates in the Gulf of Kachchh, state of Gujarat, on the western coast of India. A total of three species of the family Vermetidae were recorded during the study and their substrate preferences identified.     

Hindlimb adaptations in Ourayia and Chipetaia, relatively large-bodied omomyine primates from the Middle Eocene of Utah

North American omomyids represent a tremendous Eocene radiation of primates exhibiting a wide range of body sizes and dietary patterns. Despite this adaptive diversity, relatively little is known of the postcranial specializations of the group. Here we describe hindlimb and foot bones of Ourayia uintensis and Chipetaia lamporea that were recovered from the Uinta B member (early Uintan Land Mammal Age), Uinta Formation, Utah. These specimens provide insights into the evolution of postcranial adaptations across different body sizes and dietary guilds within the Eocene primate radiation. Body mass estimates based on talar measurements indicate that Ourayia uintensis and Chipetaia lamporea weighed about 1,500-2,000 g and 500-700 g, respectively. Skeletal elements recovered for Ourayia include the talus, navicular, entocuneiform, first metatarsal, and proximal tibia; bones of Chipetaia include the talus, navicular, entocuneiform, and proximal femur. Both genera had opposable grasping big toes, as indicated by the saddle-shaped joint between the entocuneiform and first metatarsal. Both taxa were arboreal leapers, as indicated by a consistent assemblage of characters in all represented bones, most notably the somewhat elongated naviculars, the high and distinct trochlear crests of the talus, the posteriorly oriented tibial plateau (Ourayia), and the cylindrical head of the femur (Chipetaia). The closest resemblances to Ourayia and Chipetaia are found among the Bridger omomyines, Omomys and Hemiacodon. The results of our comparisons suggest that the later, larger, more herbivorous omomyines from Utah retained a skeletal structure characteristic of earlier, smaller North American omomyids.     

The oldest mammals from Antarctica,early Eocene of the La Meseta Formation,Seymour Island

New fossil mammals found at the base of Acantilados II Allomember of the La Meseta Formation, from the early Eocene (Ypresian) of Seymour Island, represent the oldest evidence of this group in Antarctica. Two specimens are here described; the first belongs to a talonid portion of a lower right molar assigned to the sparnotheriodontid litoptern Notiolofos sp. cf. N. arquinotiensis. Sparnotheriodontid were medium‐ to large‐sized ungulates, with a wide distribution in the Eocene of South America and Antarctica. The second specimen is an intermediate phalanx referred to an indeterminate Eutheria, probably a South American native ungulate. These Antarctic findings in sediments of 55.3 Ma query the minimum age needed for terrestrial mammals to spread from South America to Antarctica, which should have occurred before the final break‐up of Gondwana. This event involves the disappearance of the land bridge formed by the Weddellian Isthmus, which connected West Antarctica and southern South America from the Late Cretaceous until sometime in the earliest Palaeogene.     

Early Eocene Primates from Gujarat, India

The oldest euprimates known from India come from the Early Eocene Cambay Formation at Vastan Mine in Gujarat. An Ypresian (early Cuisian) age of ∼53 Ma (based on foraminifera) indicates that these primates were roughly contemporary with, or perhaps predated, the India-Asia collision. Here we present new euprimate fossils from Vastan Mine, including teeth, jaws, and referred postcrania of the adapoids Marcgodinotius indicus and Asiadapis cambayensis. They are placed in the new subfamily Asiadapinae (family Notharctidae), which is most similar to primitive European Cercamoniinae such as Donrussellia and Protoadapis. Asiadapines were small primates in the size range of extant smaller bushbabies. Despite their generally very plesiomorphic morphology, asiadapines also share a few derived dental traits with sivaladapids, suggesting a possible relationship to these endemic Asian adapoids. In addition to the adapoids, a new species of the omomyid Vastanomys is described. Euprimate postcrania described include humeri, radii, femora, calcanei, and tali, most of which show typical notharctid features and are probably attributable to asiadapines. Anatomical features of the limb elements indicate that they represent active arboreal quadrupedal primates. At least one calcaneus is proximally shorter and distally longer than the others, resembling eosimiids in this regard, a relationship that, if confirmed, would also suggest an Asian or southeast Asian faunal connection. Isolated teeth from Vastan Mine recently attributed to a new eosimiid, Anthrasimias gujaratensis, appear to provide that confirmation. However, their attribution to Eosimiidae is equivocal. They are similar to teeth here tentatively referred to Marcgodinotius, hence A. gujaratensis may be a junior synonym of M. indicus. Corroboration of eosimiids at Vastan requires more compelling evidence. Although definitive conclusions are premature, available evidence suggests that the Vastan adapoids, at least, were derived from western European stock that reached India near the Paleocene-Eocene boundary.     

Early Eocene macroflora and associated palynofossils from the Cambay Shale Formation,western India: Phytogeographic and palaeoclimatic implications

A diverse assemblage of plant macrofossils and the associated representative palynofloral elements are documented from the early Eocene subsurface beds of the Cambay Shale Formation exposed in an open cast lignite mine at Vastan Village in the Surat District, western India. The Vastan mine succession is cyclic, each cycle representing a transgressive burial event terminating in the low energy lagoonal conditions. The higher energy cycle begins with sandy lenses having rich biotic remains, followed by mudstones and molluscan shell beds and ends with lignite seams. The dominantly muddy facies and the associated biota demonstrate predominantly low energy near shore or coastal plain depositional setting with conditions varying from dominantly marine (shallow) through brackish to fresh water. The Vastan mine is a well dated fossil locality with a rich and diverse biota of mammals, birds, snakes, lizards, fish, insects, molluscs, foraminifers, dinoflagellates, and plants. The plants comprise leaf and fruit impressions, seeds, fruits, wood fragments, mangrove rooting structures, fungal thalli and spores, pteridophytic spores, and angiosperm pollen grains. Thirteen macrofossil species, including several morphotaxa, are represented by the families Calophyllaceae, Rutaceae, Anacardiaceae, Rubiaceae, Combretaceae, Lythraceae, Sapindaceae, Malvaceae, and Ebenaceae. The palynological assemblage representing fourteen taxa includes the new species, Notothyrites undulatus, Callimothallus semicircularis, and Carallioipollenites integerrimoides. Habitat and distribution of modern taxa comparable with the fossil assemblage from Vastan suggest a terrestrial lowland environment. The macrofossil taxa are indicative of mesophytic, mixed forest growing under tropical to subtropical climate with sufficient humidity. The occurrence of dipterocarp elements along with taxa such as Swintonia, Pterospermum and Diospyros, etc. seems to suggest the presence of a tropical rain forest in the vicinity of Vastan.     

Intraspecific variation of Eolophiodon laboriense,a basal Lophiodontidae (Mammalia,Perissodactyla) from the early Eocene of Southern France

The Lophiodontidae is an emblematic and well-documented Eocene family of perissodactyls from Western Europe. However, after more than a century and a half of studies, lophiodontids still display a complex systematics associated with blurry intraspecific variation and a poorly known early radiation. The locality of La Borie, located near the city of Toulouse, France, has yielded numerous remains of Eolophiodon laboriense. This abundance of remains allows for the first time the study of the intraspecific variation of a basal lophiodontid. The variation has been investigated for dental and cranio-mandibular characters, notably dental polymorphism, size variation and sexual dimorphism. The intraspecific variation of E. laboriense is high with more than 20 polymorphic characters of the dentition, including many additional crests and conules. This dental polymorphism is similar to the one observed in the Bartonian lophiodontid Lophiodon lautricense. E. laboriense also displays an important degree of sexual dimorphism, with male specimens having broader and longer mandibles with larger canines than females. Despite this high intraspecific variation, the low size variation of teeth and the consistency of diagnostic characters strengthen the validity of the genus Eolophiodon and does not impact the previous lophiodontid phylogeny.     

A new genus and species of darner dragonfl y (Aeshnidae: Odonata) from the lower Eocene of Laguna del Hunco,Patagonia, Argentina

A new genus of Aeshnidae, Huncoaeshna n. gen., based on Huncoaeshna corrugata n. gen., n. sp., is erected from Laguna del Hunco (Ypresian) in Patagonia Argentina. The specimen presents a special kind of preservation with the middle part of the wing wrinkled. The presence of only two fossil specimens of Aeshnidae in South America is surely due to the lack of paleontomologists and collections of fossil insects in the subcontinent.     

A new rauisuchian archosaur from the Middle Triassic of India

Abstract:  Yarasuchus deccanensis gen. and sp. nov. is a new addition to the poorly known Middle Triassic terrestrial vertebrates. It is described on the basis of the fossil material of at least two incompletely preserved individuals and several isolated bones. It was a long-necked, gracile animal and might have had a facultatively bipedal gait . Apart from its characteristic elongate cervical vertebrae, other osteological features of Y. deccanensis resemble those of Prestosuchus, Saurosuchus, Ticinosuchus and ' Mandasuchus '. It is therefore referred to the Rauisuchian family Prestosuchidae despite the debated validity of this taxon.     

Bony‐toothed birds (Aves: Pelagornithidae) from the Middle Eocene of Belgium

Abstract: We describe well‐preserved remains of the Pelagornithidae (bony‐toothed birds) from the middle Eocene of Belgium, including a sternum, pectoral girdle bones and humeri of a single individual. The specimens are tentatively assigned to Macrodontopteryx oweni Harrison and Walker, 1976 , which has so far only been known from the holotype skull and a referred proximal ulna. Another species, about two times larger, is represented by an incomplete humerus and tentatively identified as Dasornis emuinus ( Bowerbank, 1854 ). The fossils provide critical new data on the osteology of the pectoral girdle of bony‐toothed birds. For the first time, the sternum of one of the smaller species is preserved, and this bone exhibits a more plesiomorphic morphology than the recently described sternum of the giant Miocene taxon Pelagornis. The coracoid resembles that of the Diomedeidae (albatrosses) in overall morphology, but because bony‐toothed birds lack apomorphies of the Procellariiformes, the similarities are almost certainly owing to convergence. Bony‐toothed birds were often compared with the ‘Pelecaniformes’ by previous authors, who especially made comparisons with the Sulidae (gannets and boobies). However, the coracoid distinctly differs from that of extant ‘pelecaniform’ birds, and the plesiomorphic presence of a foramen nervi supracoracoidei as well as the absence of a well‐delimited articulation facet for the furcula supports a position outside the Suloidea, the clade to which the Sulidae belong.     

Early Eocene lagomorph (Mammalia) from Western India and the early diversification of Lagomorpha

We report the oldest known record of Lagomorpha, based on distinctive, small ankle bones (calcaneus and talus) from Early Eocene deposits (Middle Ypresian equivalent, ca 53 Myr ago) of Gujarat, west-central India. The fossils predate the oldest previously known crown lagomorphs by several million years and extend the record of lagomorphs on the Indian subcontinent by 35 Myr. The bones show a mosaic of derived cursorial adaptations found in gracile Leporidae (rabbits and hares) and primitive traits characteristic of extant Ochotonidae (pikas) and more robust leporids. Together with gracile and robust calcanei from the Middle Eocene of Shanghuang, China, also reported here, the Indian fossils suggest that diversification within crown Lagomorpha and possibly divergence of the family Leporidae were already underway in the Early Eocene.     

Eocene Bird and Mammal Tracks from the Karaj Formation,Tarom Mountains,Northwestern Iran

A long continuous trackway of a shorebird (cf. Charadriipeda) and a suite of possible Pantodonta (?Corophydon) or Dinocerata tracks are reported from the Eocene Karaj Formation. Fossil footprints are still rare in Iran, described only from two Jurassic and three Tertiary sites. Thus the current record, the third in the Teritary, adds to a sparse record and increases the diversity of known track types.