Astragali of Pakicetidae and other early-to-middle Eocene archaeocetes (Mammalia,Cetacea) of Pakistan: locomotion and habitat in the initial stages of whale evolution

Richard Dehm and colleagues of the Bayerische Staatssammlung für Paläontologie und Geologie in Munich made an important collection of early-to-middle Eocene mammals at Ganda Kas in Pakistan during the winter of 1955/56. The genera and species Ichthyolestes pinfoldi and Gandakasia potens were named from this collection. Both are now recognized as early and primitive archaeocete cetaceans. In addition, Dehm’s group collected 16 complete or partial astragali of archaeocetes that were misidentified as artiodactyls. These bring the total number of archaeocete astragali known from Ganda Kas to 28. They separate clearly into four species distinguished by size: from smallest to largest Ichthyolestes pinfoldi Dehm and Oettingen-Spielberg, Pakicetus attocki (West), Gandakasia potens Dehm and Oettingen-Spielberg, and Ambulocetus natans Thewissen et al. Ganda Kas artiodactyls are smaller and rare in comparison. Ichthyolestes and Pakicetus are pakicetid archaeocetes, Gandakasia is presently indeterminate to family, and Ambulocetus is an ambulocetid. Tooth size and astragalus size are highly correlated, corroborating reference of astragali to the first three archaeocete taxa based on teeth. Multivariate morphometric comparison (Auto3Dgm) shows that pakicetid astragali overlap almost completely in shape with those of early artiodactyls. Middle Eocene protocetid astragali are divergent from both. Retention of an astragalus indistinguishable from that of artiodactyls shows that pakicetids are closely related to artiodactyls phylogenetically, but does not make Ichthyolestes and Pakicetus terrestrial or cursorial. Other skeletal elements and bone microstructure indicate that pakicetids were semiaquatic like later protocetids. Tropical riverine and marginal marine facies of the Kuldana Formation are likely habitats for initial stages of the transition from terrestrial artiodactyls to semiaquatic and fully aquatic archaeocetes.     

Bone-Breaking Bite Force of Basilosaurus isis (Mammalia,Cetacea) from the Late Eocene of Egypt Estimated by Finite Element Analysis

Bite marks suggest that the late Eocence archaeocete whale Basilosaurus isis (Birket Qarun Formation, Egypt) fed upon juveniles of the contemporary basilosaurid Dorudon atrox. Finite element analysis (FEA) of a nearly complete adult cranium of B. isis enables estimates of its bite force and tests the animal’s capabilities for crushing bone. Two loadcases reflect different biting scenarios: 1) an intitial closing phase, with all adductors active and a full condylar reaction force; and 2) a shearing phase, with the posterior temporalis active and minimized condylar force. The latter is considered probable when the jaws were nearly closed because the preserved jaws do not articulate as the molariform teeth come into occulusion. Reaction forces with all muscles active indicate that B. isis maintained relatively greater bite force anteriorly than seen in large crocodilians, and exerted a maximum bite force of at least 16,400 N at its upper P³. Under the shearing scenario with minimized condylar forces, tooth reaction forces could exceed 20,000 N despite lower magnitudes of muscle force. These bite forces at the teeth are consistent with bone indentations on Dorudon crania, reatract-and-shear hypotheses of Basilosaurus bite function, and seizure of prey by anterior teeth as proposed for other archaeocetes. The whale’s bite forces match those estimated for pliosaurus when skull lengths are equalized, suggesting similar tradeoffs of bite function and hydrodynamics. Reaction forces in B. isis were lower than maxima estimated for large crocodylians and carnivorous dinosaurs. However, comparison of force estimates from FEA and regression data indicate that B. isis exerted the largest bite forces yet estimated for any mammal, and greater force than expected from its skull width. Cephalic feeding biomechanics of Basilosaurus isis are thus consistent with habitual predation.     

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.     

A new basilosaurid (Cetacea, Pelagiceti) from the Late Eocene to Early Oligocene Otuma Formation of Peru

Cynthiacetus peruvianus nov. sp. is a new basilosaurid species, from Late Eocene to Early Oligocene Otuma Formation of Peru. It is the first described archaeocete in South-America and is represented by a sub-complete skeleton. C. peruvianus differs from C. maxwelli (middle to Late Eocene of Egypt and United States) principally in having one cuspid less on both mesial and distal sides of p3 and p4. Cynthiacetus is among the largest basilosaurids. Its more characteristic features are located on its postcranial skeleton: large vertebrarterial foramina on cervical vertebrae and absence of ventral expansion of the transverse process on C3-C5. Besides, C. peruvianus presents the greatest number of thoracic vertebrae (20) and ribs observed in Cetacea and the first thoracics have an almost vertical neural spine. A preliminary parsimony analysis establishes the monophyly of the Basilosauridae on the basis of three unambiguous cranial synapomorphies. However, within the Basilosauridae, the most diagnostic characters are observed on the postcranial skeleton.     

Gobiatherium (Mammalia: Dinocerata) from the Middle Eocene of Asia: Taxonomy and biochronological significance

Gobiatherium Osborn &; Granger 1932 is a highly distinctive genus of dinoceratan known from Eocene strata in Kyrgyzstan, Kazakstan, and Nei Mongol and Henan, People’s Republic of China. Autapomorphies ofGobiatherium among uintatheriids are: skull long, narrow and shallow with a flat sagittal region; nasals strongly arched with bony septum connecting their anterior ends to the premaxillae; zygomatic arches broad and flare outward in region of glenoid fossae; lower jaw long and shallow with tall ascending ramus and without an inframandibular flange; no upper canines; lower canines greatly reduced; P2 has prominent cingular cusp on its anterolabial corner; metacarpals relatively long and slender. Two species ofGobiatherium are valid, the type speciesG. mirificum Osborn &; Granger 1932 (=G.? major Qi 1987) and the much smaller and poorly knownG. minutum Cheng &; Ma 1990. G.?monolobotum Qi 1987 is a junior subjective synonym of the coryphodontid pantodontEudinoceras mongoliensis Osborn 1924. The distribution ofGobiatherium supports recognition of an Arshantan land-mammal ‘age’ (LMA) older than the Irdinmanhan LMA. The Arshantan LMA is redefined as the time interval between the first appearance datum (FAD) of the coryphodontid pantodontEudinoceras and the FAD of the brontotheriid perissodactylProtitan. Gobiatherium is one of several index taxa that allow mammalian assemblages of Arshantan age to be identified in China, Mongolia, Kazakstan and Kyrgyzstan.     

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.     

A new genus of rodents (Remyidae, Mammalia) from the Iberian Eocene

Abstract:  In this article, a new genus, Frontanyamys , is defined based on the new species F. russelli . The genus is recorded from the lower Upper Eocene (Bartonian) beds AT Sant Jaume de Frontanya (NE Spain). This genus shows clear affinities with the previously described genera Zamoramys, Remys and Pairomys. They are therefore assembled in the family Remyidae (new rank). The remyids are characterised by the precocious development of high-crowned molars and retain a morphologically primitive dental pattern. The oldest remyids are found in the middle Eocene of Spain ( Zamoramys ) and are probably derived from a morphologically primitive protrogomorph rodent such as Corbarimys paisi . The last representatives of the family such as Remys and Pairomys developed a fully lophodont dental pattern.     

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).     

Anthracobunids from the Middle Eocene of India and Pakistan Are Stem Perissodactyls

Anthracobunidae is an Eocene family of large mammals from south Asia that is commonly considered to be part of the radiation that gave rise to elephants (proboscideans) and sea cows (sirenians). We describe a new collection of anthracobunid fossils from Middle Eocene rocks of Indo-Pakistan that more than doubles the number of known anthracobunid fossils and challenges their putative relationships, instead implying that they are stem perissodactyls. Cranial, dental, and postcranial elements allow a revision of species and the recognition of a new anthracobunid genus. Analyses of stable isotopes and long bone geometry together suggest that most anthracobunids fed on land, but spent a considerable amount of time near water. This new evidence expands our understanding of stem perissodactyl diversity and sheds new light on perissodactyl origins.     

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.     

Comparative morphology and evolution of the otic region in toothed whales (Cetacea, Mammalia)

The otic region in the skull of archeocetes and odontocetes is compared and interpreted with special emphasis on the morphology and suspension of the ear bones. In archeocetes, the periotic was obviously separate from the mastoid but still integrated within the skull via a long anterior and posterior process. The rotation of the cochlear part of the periotic was already obvious. The tympanic bone was attached to a decreasing number of neighboring elements, with the periotic becoming more and more important in the later archeocetes. The accessory air sacs of the tympanic cavity had invaded some of the adjacent skeletal elements and attained a moderate-to-remarkable extension. In the evolution of the odontocetes, the periotic and tympanic were successively uncoupled from the skull and combined to a new morphological and functional unit (tympanoperiotic complex). This uncoupling was mainly achieved by shortening the periotical processes and simultaneously extending the tympanic air sacs. For functional reasons, however, the periotic (posterior process) stayed in immediate contact with the mastoid, the latter remaining in the lateral wall of skull. In advanced marine dolphins, the bony sheaths of the accessory air sacs are largely reduced, presumably because of volume fluctuations in the tympanic cavity during diving. The perfect uncoupling of the ear bones from the skull obviously was an essential prerequisite for directional hearing, for effective ultrasound orientation and communication, and finally, for the striking development of the dolphin brain.     

New ceratomorph perissodactyls (Mammalia) from the Middle and Late Eocene of Mongolia: their implications for phylogeny and dating

Eight species rank taxa arc described from Eocene sites in the Gobi Desert, Mongolia. Irdinolophus gen. now is erected for Desmalotherium mongoliense Osborn and for Irdinolophus? tuiensis sp. nov. The genus is placed at the stem of the family Depcrctellidae. Intraspecific variation in lekolophus magnus is documented for the first time; Pachylophus is synonymized with leleoloplms; and Deperetella cf.birmanica is recorded far north of its previously known range in Burma. The first known lower teeth of Colodon inceptus support referral ofDesmalotherium Jissum to Colodon. The stratigraphic range of the hyracodontid Ardynia is extended back from the late Eocene to the early middle Eocene. A range of primitive tapiroids and rhinocerotoids arc analysed cladistically to resolve the problematic affinities of the extinct families Dc-peretellidae and Rhodopagidae. The Depcrctellidae are shown to belong to a monophyletic superfamily Tapiroidca, whilst the Rhodopagidae belongs to the Rhinocerotoidea, but is distinct from the Hyracodontidae. Veragromovia, previously synonymized with the tapiroid Helaleles, is resurrected as a valid genus within the Rhodopagidae. Tsagan Khutcl and other sites in the Kholbolchi formation may be older than Irdinmanhan faunas in Asia, equaling with those from the Arshanto Formation of China of probable early Lutetian or latest Ypresian age. The Irdinmanhan Mergen fauna may be slightly older than that from Irdin Manha, because of the primitive aspect of its Lophialetes.     

First record of Entelodontidae (Mammalia,Artiodactyla) from the late Eocene of Southeast Asia

We describe two entelodontid upper premolars that were recovered from the late Eocene of the Krabi coal mine in southern Thailand. The size and morphology of the material suggest that it can be referred to Entelodon aff. E. gobiensis, a species known from the late Eocene to the early Oligocene of northern Asia and southern China. The Thai material documents for the first time the southernmost occurrence of entelodontids in Asia during the Paleogene and also suggests that Eocene Southeast Asian mammal localities might potentially yield further entelodontid remains mostly associated with selenodont ungulates.     

Species tree of a recent radiation: the subfamily Delphininae (Cetacea, Mammalia)

Lineages undergoing rapid radiations provide exceptional opportunities for studying speciation and adaptation, but also represent a challenge for molecular systematics because retention of ancestral polymorphisms and the occurrence of hybridization can obscure relationships among lineages. Dolphins in the subfamily Delphininae are one such case. Non-monophyly, rapid speciation events, and discordance between morphological and molecular characters have made the inference of phylogenetic relationships within this subfamily very difficult. Here we approach this problem by applying multiple methods intended to estimate species trees using a multi-gene dataset for the Delphininae (Sousa, Sotalia, Stenella, Tursiops, Delphinus and Lagenodelphis). Incongruent gene trees obtained indicate that incomplete lineage sorting and possibly hybridization are confounding the inference of species history in this group. Nonetheless, using coalescent-based methods, we have been able to extract an underlying species-tree signal from divergent histories of independent genes. This is the first time a molecular study provides support for such relationships. This study further illustrates how methods of species-tree inference can be very sensitive both to the characteristics of the dataset and the evolutionary processes affecting the evolution of the group under study.     

Pakicetus inachus and the origin of whales and dolphins (Mammalia: Cetacea)

The present paper is concerned with the comparative morphology of the archeocete and odontocete skull. Among the archeocetes, the recently described lower Eocene Pakicetus inachus obviously represents an early stage of adaptation to aquatic life. The morphology of the incomplete cranial remains, however, gives no evidence that Pakicetus was an amphibious intermediate stage. The evolution of advanced archeocetes and odontocetes is characterized by the successive acquirement of new morphological devices related to the emission and perception of ultrasound under water. The formation of a sonar system in odontocetes obviously not only helped to compensate for the loss of the peripheral olfactory system but moreover was a substantial factor in the evolution of the exceptional dolphin brain.     

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.