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.     

Discovery of a highly-specialized plesiadapiform primate in the early-middle Eocene of northwestern Africa

In this paper we report the first occurrence of an endemic African plesiadapiform primate from the early-middle Eocene locality of Glib Zegdou (Hammada du Dra, Algeria). Dralestes (new genus) is a very specialized taxon, and its closest known relative is the enigmatic and controversial genus Azibius from Gour Lazib (Hammada du Dra). We group both together as the Azibiidae (new rank). Dralestes provides the first evidence of the upper dentition in this group. Some critical dental characters, such as a postprotocingulum on upper teeth, consistently reveal a primate status for the azibiids. Dralestes exhibits, however, a very unusual configuration of the upper molars by the enlarged parastyle, the lack of a metaconule, and the ectoloph structure (preparacrista, centrocrista and postmetacrista are aligned in a high blade-like structure). The apparent dental specializations of both lower premolars and molars of azibiids (exaenodonty, large P(4) bearing sharp apical cusps, and M(1) having a highly elongated trigonid) point to potential relationships with Chronolestes and carpolestid plesiadapiforms. A phylogenetic analysis, performed on 55 dental characters scored for 19 primate genera, clarifies the euprimate status of Altiatlasius, and thus indicates that azibiids are the only known plesiadapiforms from Africa. Azibiids are the sister group of the clade carpolestids/Chronolestes in the superfamily Plesiadapoidea. However, the azibiids differ fundamentally from carpolestids by the combined lack of a centroconule and multiple buccal cusps on P(4). The exact position of both Chronolestes and azibiids in the plesiadapoids appears difficult to resolve. A basal position of Chronolestes in this superfamily cannot be ruled out because it exhibits a simple morphology of I(1) and no conule on P(3). Considering this ad hoc hypothesis, azibiids are found to lie outside a clade including carpolestids/plesiadapids/saxonellids, and they are the sister group to Chronolestes. The clade including the carpolestid, saxonellid, and plesiadapid families is characterized by the occurrence of a centroconule on P(3-4). The lack of this trait in Dralestes and Chronolestes could mean that azibiids are basal plesiadapoids that diverged before the evolution of the common ancestor of the three derived plesiadapoid families, i.e. at least around the Paleocene-Eocene boundary or more probably during the Paleocene. The report of the first offshoot in Africa of plesiadapoids enhances the role of Africa in the early primate radiation.     

Amphibians and squamates from the middle Eocene of Namibia,with comments on pre-Miocene anurans from Africa

Three middle Eocene localities (Silica North, Silica South, Black Crow) recently discovered in Namibia have produced terrestrial faunas that rank among the few known from the period of insulation of Africa (Aptian-early Miocene). Collectively, the three localities have yielded anuran amphibians (one pipid frog, the earliest assemblage [three taxa] of ranoid frogs in Africa, one indeterminate family) and squamate reptiles (an amphisbaenian ‘lizard’, a snake that likely represents a colubroid, and two indeterminate ‘lizards’). These Eocene faunas suggest that ranoids, colubroids and African pipids are autochthonous to Africa. However, whereas pipids are vicariants inherited from West Gondwana, ranoids and colubroids (if really autochthonous) originated in Africa from unknown stems. Silica North and Silica South correspond to aquatic environments, permanent fresh water being present in the first locality; the environment of Black Crow was drier.     

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.     

Functional differentiation of trailing and leading forelimbs during locomotion on the ground and on a horizontal branch in the European red squirrel (Sciurus vulgaris, Rodentia)

Mammalian locomotion is characterized by the frequent use of in-phase gaits in which the footfalls of the left and right fore- or hindlimbs are unevenly spaced in time. Although previous studies have identified a functional differentiation between the first limb (trailing limb) and the second limb (leading limb) to touch the ground during terrestrial locomotion, the influence of a horizontal branch on limb function has never been explored. To determine the functional differences between trailing and leading forelimbs during locomotion on the ground and on a horizontal branch, X-ray motion analysis and force measurements were carried out in two European red squirrels (Sciurus vulgaris, Rodentia). The differences observed between trailing and leading forelimbs were minimal during terrestrial locomotion, where both limbs fulfill two functions and go through a shock-absorbing phase followed by a generating phase. During locomotion on a horizontal branch, European red squirrels reduce speed and all substrate reaction forces transmitted may be due to the reduction of vertical oscillation of the center of mass. Further adjustments during locomotion on a horizontal branch differ significantly between trailing and leading forelimbs and include limb flexion, lead intervals, limb protraction and vertical displacement of the scapular pivot. Consequently, trailing and leading forelimbs perform different functions. Trailing forelimbs function primarily as shock-absorbing elements, whereas leading forelimbs are characterized by a high level of stiffness. This functional differentiation indicates that European red squirrels ‘test’ the substrate for stability with the trailing forelimb, while the leading forelimb responds to or counteracts swinging or snapping branches.     

New marine mammal faunas (Cetacea and Sirenia) and sea level change in the Samlat Formation,Upper Eocene,near Ad-Dakhla in southwestern Morocco

The Samlat Formation is well exposed in coastal sections bordering the Atlantic Ocean south of Ad-Dakhla in southwestern Morocco. Here some 22 m of rhythmically-bedded, chert-rich, marine siltstones and marls are overlain by 1–1.5 m of vertebrate-bearing microconglomeratic sandstone, another 4–8 m of rhythmically-bedded siltstone and marl, and finally a second 3–6 m unit of vertebrate-bearing muddy sandstone. The microconglomeratic and muddy sandstones represent low sea stands in what is otherwise a deeper water sequence. Cetacean skeletons are rare but cetacean vertebrae are common in the lower sandstone (bed B1), where many show the effects of reworking. The cetaceans in bed B1represent a minimum of five species, from smallest to largest: cf. Saghacetus sp., cf. Stromerius sp., Dorudon atrox, cf. Dorudon sp., and Basilosaurus isis. Bed B1 yields rib fragments that may represent sirenians, but sirenians, if present, are rare. The only identifiable cetacean found in the upper sandstone (bed B2) is Basilosaurus sp. Dugongid sirenians identified as cf. Eosiren sp. are the most common mammal in bed B2. We interpret co-occurrence of the typically Early Priabonian species Dorudon atrox and Basilosaurus isis with smaller species more like Middle Priabonian genera Saghacetus osiris and Stromerius nidensis to indicate that bed B1 was deposited during low sea stand Pr-2 between the Early and Middle Priabonian (between the early and middle Late Eocene). Bed B2 is separated from B1 by an interval of deeper water sediment accumulation. Bed B2 could represent a later phase of Pr-2 or a subsequent Priabonian low sea stand (possibly Pr-3).     

Basilotritus (Cetacea: Pelagiceti) from the Eocene of Nagornoye (Ukraine): New data on anatomy,ontogeny and feeding of early basilosaurids

A new specimen of early basilosaurid, now identified as Basilotritus sp., comes from the late Middle Eocene of Ukraine. It has basilosaurid-type cheek teeth with cinguli, similar to those of Zygorhiza, and roots resembling those of Georgiacetus vogtlensis and early Neoceti; an unusual feature of these teeth is the presence of accessory denticles of the second order located on the crown denticles. The postcranial anatomy shows a mixture of primitive and advanced basilosaurid traits. The phylogenetic position of the genus Basilotritus is confirmed to be near the base of Basilosauridae, between Supayacetus and Zygorhiza. The ontogeny of the specimen from Nagornoye is characterized by large body size, slow skeletal maturation and intensive pachyosteosclerosis that are interpreted as neotenic development. Rapid tooth wear with strong apical abrasion is the result of specialized diet, possibly feeding on sharks.     

张和兽(Zhangheotherium)的齿列和前肢形态

张和兽（Zhangheotherium）是目前唯一保存完整骨架的对齿兽类。其西列特征表明它属于兽科（Spalacotheriidae）,该科为原始兽类中主要繁衍于北方大陆中生代后期一个咀嚼方式以上下日齿对咬为主的单系类群。张和兽骨骼形态显示其前肢姿势处于外展趴卧（sprawling）与内收直立（parasagittal）之间,兼有树上和地面活动的能力,是原始兽类中趋于以地面活动为主的成员。附录列举了哺乳类的主要齿列和骨骼性状序列及分布矩阵。