New remains of Chambius kasserinensis from the Eocene of Tunisia and evaluation of proposed affinities for Macroscelidea (Mammalia,Afrotheria)

Abstract

Chambius kasserinensis from the late Early or early Middle Eocene Chambi locality, central Tunisia, is undoubtedly the oldest known macroscelidid and possibly the basalmost representative of the order Macroscelidea. Hence, since its discovery in 1986, Chambius has played a key role in analyses focusing on afrotherian and eutherian phylogeny; for instance, as early as 1995, Butler’s review of fossil macroscelideans highlighted the central position of Chambius in the origin of the order. Despite this, Chambius remained poorly known until recently. Here based on new mandibular fragments, well-preserved upper molars and CT scan analysis of the holotype maxilla, Chambius is revised. Its dentition is first described in detail, providing a precise characterization of the genus. Chambius is notably defined by a submolariform P₄ with a three-cusped talonid, a reduced talonid on M₂, and a prominent metaconule on M^1?2. Interestingly, the two transverse lophs of the upper molars are basically formed by preconulecristae, evoking the recently defined peculiar bilophodonty of paenungulates. Comparisons with other Paleogene and modern macroscelidids, European Louisinidae, and North American Apheliscidae are also made, allowing the various hypotheses about the origin and early evolution of macroscelidids to be reviewed.     

Relative importance of habitat connectivity in shaping the genetic profiles of two southern African elephant‐shrews

Aim When interpreting genetic patterns across a landscape it is surprisingly difficult to disentangle the effects of landscape connectivity from those of species biology. Here, the spatial distributions of genetic variation of two sympatric elephant‐shrew species, the western rock elephant‐shrew (Elephantulus rupestris) and the round‐eared elephant‐shrew (Macroscelides proboscideus), are determined and compared. We selected these species because they have similar biologies but differ markedly in habitat use, the rationale being that differences in their genetic structure should be a result largely of landscape variables directly or indirectly affecting dispersal rather than of the biology of the species. Location South Africa and Namibia. Methods Mitochondrial sequence data (control region and cytochrome b) were used to describe the phylogeographic structure of these elephant‐shrew species across their distribution. To determine whether genetic variation is significantly structured, spatial analyses of molecular variation were performed. Isolation‐by‐distance versus alternative patterns of genetic structure was investigated using a Mantel test. Results Our analyses indicated an overall structured genetic profile for E. rupestris, a species closely associated with rocky outcrops. This was in contrast to a pattern mostly of isolation‐by‐distance across the distribution of M. proboscideus, a species found on gravel plains. Main conclusions Specific landscape features will differentially affect gene flow (both historical and current), and therefore also the spatial genetic structure, of species with markedly different habitat requirements. The genetic profiles for the two species included here support predictions based on the connectivity of their respective occupied habitats. The results also support the more general prediction that species with a naturally clustered distribution (such as E. rupestris) should have a more structured genetic pattern than those having a more continuous distribution (M. proboscideus).     

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 Fish Assemblage from the Middle Eocene from Libya (Dur At-Talah) and the Earliest Record of Modern African Fish Genera

In the early nineteen sixties, Arambourg and Magnier found some freshwater fish (i.e., Polypterus sp., Siluriformes indet. and Lates sp.) mixed with marine members in an Eocene vertebrate assemblage at Gebel Coquin, in the southern Libyan Desert. This locality, aged ca 37–39Ma and now known under the name of Dur At-Talah, has been recently excavated. A new fish assemblage, mostly composed of teeth, was collected by the Mission Paléontologique Franco-Libyenne. In this paper, we describe freshwater fish members including a dipnoan (Protopterus sp.), and several actinopterygians: bichir (Polypterus sp.), aba fish (Gymnarchus sp.), several catfishes (Chrysichthys sp. and a mochokid indet.), several characiforms (including the tiger fish Hydrocynus sp., and one or two alestin-like fish), and perciforms (including the snake-head fish Parachanna sp. and at least one cichlid). Together with the fossiliferous outcrops at Birket Qarun in Egypt, the Libyan site at Dur At-Talah reduces a 10-Ma chronological gap in the fossil record of African freshwater fish. Their fish assemblages overlap in their composition and thus constitute a rather homogenous, original and significant amount of new elements regarding the Paleogene African ichthyofauna. This supports the establishment of the modern African freshwater fish fauna during this time period because these sites mostly contain the earliest members known in modern genera.     

Confirmation of pleisiomorphic daily torpor in mammals: the round-eared elephant shrew Macroscelides proboscideus (Macroscelidea)

The characteristics of daily torpor were measured in the round-eared elephant shrew Macroscelides proboscideus (Macroscelidea) in response to ambient temperature and food deprivation. Elephant shrews are an ancient mammal order within a superordinal African clade including hyraxes, elephants, dugongs and the aardvark. M. proboscideus only employed torpor when deprived of food; torpor did not occur under an ad libitum diet at ambient temperatures of 10, 15 and 25?°C. Torpor bout duration ranged from <1?h to ca. 18?h. The times of entry into torpor were restricted to the scotophase, despite normothermic body temperature patterns indicating a rest phase coincident with the photophase. Full arousal was always achieved within the first 3?h of the photophase. When food deprived, the onset of the rest phase, and hence torpor, advanced with respect to the experimental photoperiod. The lowest torpor body temperature measured was 9.41?°C. Daily torpor in M. proboscideus confirms a pleisiomorphic origin of daily heterothermy. Torpor facilitates risk-averse foraging behaviour in these small omnivores by overcoming long-term energy shortfalls generated by the inherent variability of food availability in their semi-arid, El Niño-afflicted habitats.     

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.     

Evolution of the tooth enamel microstructure in the earliest proboscideans (Mammalia)

Microstructural features of the mammalian tooth enamel are rarely used to construct phylogenies, although macromorphological characters of the dentition figure prominently in phylogenetic analysis. In order to test the phylogenetic significance of the enamel microstructures, we investigate here the earliest proboscideans recently found in the Early Palaeogene of Africa (e.g. Phosphatherium , Daouitherium , Khamsaconus , and Numidotherium ). The results are discussed in the light of the recent advances concerning the intra- and interordinal relationships of the Proboscidea. We also consider other basal paenungulates such as 'anthracobunids', embrithopods, and hyraxes. The analysed microstructures suggest that the enamel ancestral morphotype of paenungulates was primitive for eutherian mammals, consisting in radial enamel. Some basal proboscideans developed decussations of prisms in Hunter-Schreger bands (HSB), as did most of the medium to large-sized mammals. More evolved proboscideans developed very complex enamel, the 3-D enamel, which represents an apomorphy for the group. The three-layered Schmelzmuster, typical of the elephantoids (3-D enamel, HSB, and radial enamel), is acquired during the late Eocene with the enigmatic ' Numidotherium ' savagei . This species is here considered as an advanced proboscidean along with Moeritherium -Deinotheriidae-Elephantiformes. The peculiar enamel of elephantoids arose step by step. Although homoplasy and mosaic evolution occur, the enamel microstructures represent an important source of new dental characters for phylogenetic reconstructions. As macromorphological characters testified, the diversity of the enamel microstructures observed in the various basal proboscideans illustrates an unexpected early diversity of the order in Africa.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 149 , 611–628.     

A New Eocene Dasypodid with Caniniforms (Mammalia,Xenarthra, Cingulata) from Northwest Argentina

The aim of this work is to present a new genus and species of Dasypodidae from the Lumbrera Formation (“lower Lumbrera”), early-middle Eocene of Salta Province, northwest Argentina. The new taxon, documented by one specimen, consists of an incomplete skull and jaw with teeth housed in their alveoli, postcranial remains, and isolated osteoderms. Lumbreratherium oblitum, gen. et sp. nov., is characterized by heterodont dentition, with a caniniform as the first tooth, diastema between caniniform and first molariform, teeth with closed roots, and a peculiar morphology of the osteoderms. A phylogenetic analysis indicates that Lumbreratherium oblitum, gen. et sp. nov., and Pucatherium parvum belong to a monophyletic clade in a basal position within the Cingulata. The singularity of the morphological characteristics of these Paleogene armadillos of northwest Argentina reinforces the hypothesis of an intertropical origin of mammal clades different from those of the Paleogene in more austral regions of Argentina.     

New remains of Egatochoerus jaegeri (Mammalia,Suoidea) from the late Eocene of Peninsular Thailand

Abstract: We report here additional remains referred to Egatochoerus jaegeri from the late Eocene locality of Ban Mark in the Krabi basin, Thailand. The new material described, comprising upper and lower cheek teeth, deciduous premolars and partial cranial remains, makes E. jaegeri the best‐documented Eocene representative of Old World Suoidea at present. Detailed study and comparison of their cheek teeth structure reveal a homogeneity of the molar crest and groove patterns of Old World and New World Palaeogene suoids.     

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 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.     

Post-Pleistocene evolution of Bornean shrews Crocidura foetida (Mammalia, Soricidae)

Fossil mandibles of the Bornean shrew Crocidura foetida recovered from excavations at the west mouth of Niah cave, Sarawak, Malaysia, show that the late Pleistocene population at this lowland location was comparable in size with the large subspecies Crocidura foetida doriae , presently occurring at inland, upland locations. Two Holocene specimens fall in the size range of the smaller lowland subspecies C. f. foetida . Comparable post-Pleistocene size-reduction is known among other mammals of Borneo, but this is the first instance of dated examples. The evolutionary trend conforms with Bergmann's 'rule' but, other than climate change, no selective agent is apparent.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 413–419.     

Crocodilian remains from the Upper Eocene of Dor-El-Talha, Libya

Study of undescribed material from Dor-El-Talha (Libya, Upper Eocene) reveals the existence of at least three different crocodilian taxa at this locality: a gavialid, a tomistomine and a mesosuchian represented by just one amphicoelous vertebra. Most of this material is fragmentary and it includes mainly fragments of snouts and lower jaws. The crocodilian fauna from this locality shows similarities with that from the contemporaneous El Fayum locality (Egypt).

Résumé

L’étude du matériel inédit provenant de Dor-El-Talha (Libye, Éocène supérieur) indique l’existence d’au moins trois taxons différents de crocodiliens : un gavialidé, un tomistomine et un mésosuchien représenté par une seule vertèbre amphicoele. La plupart de ces matériaux est fragmentaire et elle comprend principalement des fragments de museaux et mandibules. La faune crocodilienne de ce gisement montre des ressemblances avec celle provenant du gisement contemporain du Fayum (Egypte).     

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.     

Phylogeny and taxonomy of the round-eared sengis or elephant-shrews, genus Macroscelides (Mammalia, Afrotheria, Macroscelidea)

The round-eared sengis or elephant-shrews (genus Macroscelides) exhibit striking pelage variation throughout their ranges. Over ten taxonomic names have been proposed to describe this variation, but currently only two taxa are recognized (M. proboscideus proboscideus and M. p. flavicaudatus). Here, we review the taxonomic history of Macroscelides, and we use data on the geographic distribution, morphology, and mitochondrial DNA sequence to evaluate the current taxonomy. Our data support only two taxa that correspond to the currently recognized subspecies M. p. proboscideus and M. p. flavicaudatus. Mitochondrial haplotypes of these two taxa are reciprocally monophyletic with over 13% uncorrected sequence divergence between them. PCA analysis of 14 morphological characters (mostly cranial) grouped the two taxa into non-overlapping clusters, and body mass alone is a relatively reliable distinguishing character throughout much of Macroscelides range. Although fieldworkers were unable to find sympatric populations, the two taxa were found within 50 km of each other, and genetic analysis showed no evidence of gene flow. Based upon corroborating genetic data, morphological data, near sympatry with no evidence of gene flow, and differences in habitat use, we elevate these two forms to full species.     

White-toothed shrews (Mammalia, Soricomorpha, Crocidura) of coastal islands of Vietnam

New findings of the white-toothed shrews (Crocidura spp.) from offshore islands of Vietnam are reported. The species identifications have been confirmed by the analysis of complete mitochondrial cytochrome b gene (1140 bp). Crocidura phuquocensis is the only species found in the Phu Quoc Island. Crocidura fuliginosa has been recorded from two islands of the Con Dao Archipelago (Con Son and Bai Canh). The occurrence of Crocidura fuliginosa in Vietnam has been genetically confirmed for the first time. Crocidura attenuata has been collected from the Cat Ba Island for the first time, and this finding corresponds well with the proposal that the species' distribution is confined to the north and east of the Red River only.