Eocene ziphodont crocodilia from Northwestern India

The Middle Eocene (Lutetian) strata constituting the top of the Subathu Formation in Jammu and Kashmir, have yielded a large number of ziphodont crocodilians. The affinity of these serrated-toothed crocodiles has been assigned to the Eusuchian Pristichampsus rather than to the Sebecosuchia. The teeth are morphologically diverse, strongly fluted with serrations varying from fine to coarse. It is believed that the Kashmir ziphodont crocodiles are migrant descendants of Chinese forms which spread across the Indian sub-continent. Their presence in Europe may be accounted by migration along with Asiatosuchus across the Turgai Straits sometime in the Early Eocene.     

New Protocetid Whale from the Middle Eocene of Pakistan: Birth on Land,Precocial Development,and Sexual Dimorphism

Background

Protocetidae are middle Eocene (49–37 Ma) archaeocete predators ancestral to later whales. They are found in marine sedimentary rocks, but retain four legs and were not yet fully aquatic. Protocetids have been interpreted as amphibious, feeding in the sea but returning to land to rest.

Methodology/Principal Findings

Two adult skeletons of a new 2.6 meter long protocetid, Maiacetus inuus, are described from the early middle Eocene Habib Rahi Formation of Pakistan. M. inuus differs from contemporary archaic whales in having a fused mandibular symphysis, distinctive astragalus bones in the ankle, and a less hind-limb dominated postcranial skeleton. One adult skeleton is female and bears the skull and partial skeleton of a single large near-term fetus. The fetal skeleton is positioned for head-first delivery, which typifies land mammals but not extant whales, evidence that birth took place on land. The fetal skeleton has permanent first molars well mineralized, which indicates precocial development at birth. Precocial development, with attendant size and mobility, were as critical for survival of a neonate at the land-sea interface in the Eocene as they are today. The second adult skeleton is the most complete known for a protocetid. The vertebral column, preserved in articulation, has 7 cervicals, 13 thoracics, 6 lumbars, 4 sacrals, and 21 caudals. All four limbs are preserved with hands and feet. This adult is 12% larger in linear dimensions than the female skeleton, on average, has canine teeth that are 20% larger, and is interpreted as male. Moderate sexual dimorphism indicates limited male-male competition during breeding, which in turn suggests little aggregation of food or shelter in the environment inhabited by protocetids.

Conclusions/Significance

Discovery of a near-term fetus positioned for head-first delivery provides important evidence that early protocetid whales gave birth on land. This is consistent with skeletal morphology enabling Maiacetus to support its weight on land and corroborates previous ideas that protocetids were amphibious. Specimens this complete are virtual ‘Rosetta stones’ providing insight into functional capabilities and life history of extinct animals that cannot be gained any other way.     

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.     

Foraminifera-walled Schaubcylindrichnus coronus Frey and Howard, 1981, from the Middle Eocene,Kachchh, Western India

Bioclast lined tubes of Schaubcylindrichnus coronus are not much reported around the world. As of now only two reports are available. Interestingly in both these reports and in the current find, tests of foraminifers are used as tube building material. Current report of Schaubcylindrichnus is from the subtidal deposits of the Naredi Formation (Ypresian) of Kachchh Basin, Western India. As the host rock here abounds in the tests of larger benthic foraminifera, concentration of same is noticed within the tubes. The natural selection of lighter and larger calcitic grains of tests over the heavier mineral grains is the main reason for presence of the tests in tube building; moreover, presence of inner smooth wall of tube also indicates organisms’ preference for comfort within their burrow.     

Non-marine Teredolites from the Middle Eocene of southern England

Middle Eocene, non-marine sediments from southern England contain examples of Teredolites borings in two contrasting palaeoenvironmental settings, viz.: (A) as in situ borings in an allochthonous lignite in an abandoned river channel and (B) as bored logs in cross-bedded fluvial sandstones of probable point-bar origin. The lignite is 0.30 m thick, of which the upper 0.20 m is intensely bored. Rounded pebbles of ?charcoal at its base also show small borings. A log in a fluvial sandstone shows densely-packed. radial club-shaped borings. now filled with sandstone. The wood substrate has subsequently been oxidized away. To our knowledge, this is the first detailed account of Teredolites from an ancient freshwater setting.     

Early Eocene orthophragminids from the Lower Indus Basin,Pakistan, and their biostratigraphic implication

Orthophragminids are a key biostratigraphic proxy for Paleocene–Eocene sequences in western Tethys, but poorly known in eastern Tethys. The orthophragminids were previously misinterpreted as Paleocene's orbitoidiform foraminifera in the Indus Basin, Pakistan. This study focuses on detailed taxonomy and biostratigraphy of orthophragminids. Nine species/subspecies of orthophragminids are identified, including Discocyclina ranikotensis, D. archiaci bakhchisaraiensis, D. a. ex. interc. staroseliensis bakhchisaraiensis, D. dispansa hungarica, D. d. taurica, D. d. broennimanni and Orbitoclypeus schopeni ramaraoi, and two new species Discocyclina pseudoranikotensis n. sp. and Discocyclina debalensis n. sp. are described. The identified orthophragminid taxa represent the orthophragminid zones OZ2-3, corresponding to the shallow benthic zones SBZ5-7 of early Ypresian (early Eocene), which are correlated with those in western Tethys. However, we found that the first appearances of D. d. hungarica and D. d. taurica occur in the OZ3 or SBZ7 in this study instead of the OZ5 or SBZ10 as in western Tethys. The possible paleoecological setting for the occurrences of recognized species is interpreted as inner to middle shelf.     

Anatomy and development of fruits of Lauraceae from the Middle Eocene Princeton Chert

Investigations of the Middle Eocene Princeton Chert reveal evidence for the connection of lauraceous flowers to fruits through a developmental series. Youngest fruits are found with attached floral remnants. Later stages show receptacle enlargement, fruit wall thickening, and the development of abundant sclereid clusters. Mature fruits are borne on a shallow receptacle and have an endocarp palisade layer of radially elongate cells with stellate outlines, an inner mesocarp layer of radiately arranged sclereid clusters, and a fleshy outer mesocarp layer containing numerous idioblasts with contents. Each mature fruit bears a single seed retaining the outer integument with an innermost radially elongate transfusion cell layer. Mature seeds contain a cellular embryo bearing idioblasts. Fruits are distinguishable from previously described anatomically preserved fossil taxa. This study represents the only documented developmental reconstruction of fossil fruits of Lauraceae and that self-pruning evolved prior to the Eocene. Anatomical modifications over the developmental sequence indicate that different stages of maturity preserved together, may be erroneously identified as several taxa at a fossil locality. Fossil morphotypes typically underestimate species number, but this study suggests that the number of inferred species based on fruit types may be inflated for Lauraceae, potentially exaggerating the tropical interpretation of the paleoenvironment.     

Attockicetus praecursor, A New Remingtonocetid Cetacean from Marine Eocene Sediments of Pakistan

A new early-to-middle Eocene cetacean from the Kala Chitta Hills of northern Pakistan is described: Attockicetus praecursor new genus and species. It is based on fragmentary cranial material, including a rostral fragment, P3–M3, endocast, and ectotympanic. Attockicetus is the first remingtonocetid from northern Pakistan, and the oldest member of its family. Attockicetus praecursor is smaller than the species of the other remingtonocetid genera, Remingtonocetus, Andrewsiphius, and Dalanistes. It is also more primitive in the retention of large protocones on the upper molars and the anterior position of the orbit. Known material for Attockicetus is fragmentary, but the taxon is important because it extends the geographic and temporal range of remingtonocetids, is one of the few remingtonocetids in which toothcrowns are preserved, and because it is probably the most plesiomorphic remingtonocetid.     

New Lizards from the Middle Eocene Mergen Formation, Mongolian Gobi Desert

New lizard fossils are described from the middle Eocene Mergen Formation, eastern Gobi Desert in Mongolia. Some 30 isolated tooth-bearing jaws represent at least six lizard taxa. These include three new iguanians that are nonchamaeleonid acrodontans; two scincomorphs having uncertain relationships within the group; and one large anguimorph that may represent a platynotan taxon. Discovery of these fossil lizards is important for accumulating nonmammalian vertebrate data, in attempting to understand a major faunal change that took place in relation to the drastic uplifting of the Mongolian Plateau in Palaeogene time.     

New acrodont lizards (Lacertilia) from the Middle Eocene of southern Mongolia

The problematic lizard family Changjiangosauridae, representatives of which inhabited Asia in the Early Paleogene, is discussed. Six new species of this group, including Acrodontopsis robustus gen. et sp. nov., Agamimus gracilis gen. et sp. nov., Graminisaurus interruptus gen. et sp. nov., Khaichinsaurus reshetovi gen. et sp. nov., Lavatisaurus elegans gen. et sp. nov., and Lentisaurus giganteus gen. et sp. nov., from the Middle Eocene of the Khaichin Uul 2 locality (southern Gobi, Mongolia) are described. It is shown that Changjiangosauridae are probably related to the Late Cretaceous Isodontosauridae and recent Uromastycidae; independent development of a number of dental features in different lineages of Acrodonta (Iguania) is corroborated.     

Larger benthic foraminiferal assemblages and their response to Middle Eocene Climate Optimum in the Kohat Basin (Pakistan,eastern Tethys)

The Middle Eocene Climatic Optimum (MECO) at ～40 Ma is a significant global warming event associated with pronounced changes in the hydrosphere, atmosphere, and biosphere. The Kohat Formation in the Kohat Basin (eastern Tethys, Pakistan) is studied for identifying the response of larger benthic foraminifera (LBF) to MECO. The LBF assemblages in the Kohat Formation, covering from the Shallow Benthic Zones (SBZ) 15 to 17, suggest middle Lutetian to early Bartonian in age. Microfacies analyses indicate a lagoonal (inner carbonate ramp facies belt) to open marine (middle carbonate ramp facies belt) paleodepositional environment of the Kohat Formation. A distinct positive δ¹³C shift marks the stratigraphic position of the MECO in this formation. At the Peak-MECO event that is marked by the onset of the positive carbon isotope excursion (CIE), no evident compositional variation in the LBF assemblages is observed. However, significant changes in the LBF assemblages with the local first and last occurrences of some LBF genera can be observed in the Post-MECO and CIE recovery phase. These changes are verified by the sudden disappearance of Alveolina and orthophragminids and initial dominance of larger shell-size Nummulites fabianii, Heterostegina, and Linderina species accompanied by an increase in the species diversity. Here, we argue that the change in the observed LBF assemblages in the uppermost part of the Kohat Formation might be related to a larger foraminiferal turnover occurring during the Post-MECO event and corresponds to the CIE recovery phase.     

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.     

Halophilic Archaea Cultivated from Surface Sterilized Middle-Late Eocene Rock Salt Are Polyploid

Live bacteria and archaea have been isolated from several rock salt deposits of up to hundreds of millions of years of age from all around the world. A key factor affecting their longevity is the ability to keep their genomic DNA intact, for which efficient repair mechanisms are needed. Polyploid microbes are known to have an increased resistance towards mutations and DNA damage, and it has been suggested that microbes from deeply buried rock salt would carry several copies of their genomes. Here, cultivable halophilic microbes were isolated from a surface sterilized middle-late Eocene (38–41 million years ago) rock salt sample, drilled from the depth of 800 m at Yunying salt mine, China. Eight unique isolates were obtained, which represented two haloarchaeal genera, Halobacterium and Halolamina. We used real-time PCR to show that our isolates are polyploid, with genome copy numbers of 11–14 genomes per cell in exponential growth phase. The ploidy level was slightly downregulated in stationary growth phase, but the cells still had an average genome copy number of 6–8. The polyploidy of halophilic archaea living in ancient rock salt might be a factor explaining how these organisms are able to overcome the challenge of prolonged survival during their entombment.     

Middle Eocene rodents from Peruvian Amazonia reveal the pattern and timing of caviomorph origins and biogeography

The long-term isolation of South America during most of the Cenozoic produced a highly peculiar terrestrial vertebrate biota, with a wide array of mammal groups, among which caviomorph rodents and platyrrhine primates are Mid-Cenozoic immigrants. In the absence of indisputable pre-Oligocene South American rodents or primates, the mode, timing and biogeography of these extraordinary dispersals remained debated. Here, we describe South America's oldest known rodents, based on a new diverse caviomorph assemblage from the late Middle Eocene (approx. 41 Ma) of Peru, including five small rodents with three stem caviomorphs. Instead of being tied to the Eocene/Oligocene global cooling and drying episode (approx. 34 Ma), as previously considered, the arrival of caviomorphs and their initial radiation in South America probably occurred under much warmer and wetter conditions, around the Mid-Eocene Climatic Optimum. Our phylogenetic results reaffirm the African origin of South American rodents and support a trans-Atlantic dispersal of these mammals during Middle Eocene times. This discovery further extends the gap (approx. 15 Myr) between first appearances of rodents and primates in South America.     

A new Chalicothere from the Pondaung Formation (late Middle Eocene of Myanmar)

The 2002 field work campaign of the joint Myanmar–French Expedition has resulted in the discovery of new material particularly in the Bahin area (Pondaung Formation, Middle–Upper Eocene, Myanmar). The Nyaung Pin Le outcrop has yielded, among other fossil mammals, a maxillary of a primitive chalicothere, the first ever discovered in the southeastern Asia Palaeogene, which is assigned to the Eomoropidae, Eomoropus pawnyunti nov. sp. This form is distinguished from the other representatives of the family by its small size, its brachyodonty and the few derived shape of the molar ectolophs, nevertheless provided with low mesostyles. To cite this article: J.-A. Remy et al., C. R. Palevol 4 (2005).     

Paratritemnodon indicus (Creodonta: Mammalia) from the early Middle Eocene Subathu Formation,NW Himalaya,India, and the Kalakot mammalian community structure

A complete post-incisor upper dentition and a left mandible with P/3-M/3 of the proviverrine hyaenodontid (Creodonta: Mammalia)Paratritemnodon indicus, which was hitherto known by heavily worn P/3-M/3, are described from the uppermost Subathu Formation (Middle Eocene) of the Kalakot-Metka-Mohgala area, Rajauri District, Jammu and Kashmir.P. indicus is most closely related to the North American Early Eocene proviverrine,Tritemnodon; its relationships withProdissopsalis from the Middle Eocene of Europe andPropterodon from the Middle to Late Eocene of China are also close. In the Kalakot Eocene vertebrate community of the Subathu vertebrate bioprovince,P. indicus is the only land dwelling carnivorous mammal against more than 20 species of herbivorous mammals, a majority of which are larger than it. This is viewed as an imbalance in trie community and it reflects an immaturity of the ecosystem supporting the fauna. The scarcity of carnivore remains in the Kalakot mammalian fauna is not an artifact.     

Duabanga-like leaves from the Middle Eocene Princeton chert and comparative leaf histology of Lythraceae sensu lato

A permineralized lythraceous leaf type found in close association with fruits, stems, and roots of Decodon allenbyensis Cevallos-Ferriz et Stockey in the Middle Eocene Princeton chert of British Columbia, Canada, is described. Midribs have a prominent C-shaped midvein surrounded by sclerenchyma, with an adaxial epidermis of rectangular to rounded cells lacking enlarged mucilage cells. Leaves are dorsiventral, 180-270 μm thick at the lamina, with a double palisade layer. Abaxial epidermal cells have prominent papillae, and these epidermal cells can be infected by fungi, forming dark sterile stromata. Fossil leaves are similar to those of Myrtales and are compared to those of Lythraceae sensu lato. Although these leaves are thought to belong to the previously described Decodon allenbyensis found in the same chert layer, they lack the diagnostic features of extant Decodon leaves. Instead they share most anatomical similarities with Duabanga grandiflora Roxburgh ex DC Walpers (Lythraceae, subfamily Duabangoideae) including vascular tissues, palisade and spongy mesophyll, bundle fibers, and abaxial epidermal papillae. Duabanga grandiflora differs from the fossil in having mucilaginous cells and a consistently V-shaped abaxial midrib. Although anatomically similar to Duabanga, the fossil leaves are considered those of D. allenbyensis, based on association and the depositional environment prior to preservation. Recent phylogenetic analyses place Duabanga and Decodon in separate clades within Lythraceae, but relationships between these clades are not well supported, indicating that fossil leaves should provide useful anatomical characters for elucidating relationships within Lythraceae.