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A new neotenic salamander, Seminobatrachus boltyschkensis gen. et sp. nov., is described based on 14 skeletons of late Paleocene–early Eocene age preserved on drill core slabs from the Cherkassy Region, central Ukraine. The new taxon is diagnosed by the following unique combination of characters: dorsal process of premaxilla posteriorly elongate and overlaps frontal; maxilla greatly reduced in size; parietal–squamosal contact absent; vomerine tooth row long and parallel to maxillary arcade; pterygoid has long anterior process; quadrate ossified; marginal and palatal teeth pedicellate; trunk vertebrae amphicoelous, each having a subcentral keel, anterior basapophysis, and spinal nerve foramina; ribs bicipital; carpals and tarsals unossified; and phalangeal formulae of 2−2−3−2 and 2−2−3−4−2 for manus and pes, respectively. Phylogenetic analysis nests S. boltyschkensis within Urodela (i.e., crown−clade salamanders), but its exact phylogenetic position is equivocal, resolving in one of three ways: (1) in an unresolved trichotomy with Salamandra and (Ambystomatidae + (Dicamptodon + Rhyacotriton)) (results obtained in NONA v. 2.0, with the WINCLADA v. 1.00.08 interface; the parsimony ratchet (island hopper) algorithm), (2) as a sister taxon of (Salamandra + (Ambystomatidae + (Dicamptodon + Rhyacotriton))) clade (results obtained in TNT v. 1.1; the implicit enumeration search algorithm) or (3) as a sister taxon of Ambystomatidae (results obtained in PAUP v. 4.0b10; the branch−and−bound search algorithm).
Adapisoriculidae are an enigmatic group of small mammals known from the late Cretaceous of India, and from the early Palaeocene to early Eocene of Europe and Africa. Based on their primitive dental morphology, they have been classified as didelphids, nyctitheriids, leptictids, mixodectids, tupaiids, and palaeoryctids. While the latest hypothesis based on dental morphology suggests an affinity with Lipotyphla, postcranial remains indicate a close relationship with Euarchonta. Here, we present new adapisoriculid dental remains from the early Palaeocene locality of Hainin (Belgium). Adapisoriculidae are particularly abundant in Hainin, where they represent about one third of the mammalian fauna, offering new insights into both their specific and generic phylogenetic interrelationships. We describe three new species (Afrodon gheerbranti sp. nov., Bustylus folieae sp. nov. and Proremiculus lagnauxi gen. et sp. nov.) and document the previously unknown lower dentition of Bustylus marandati. The diversity of dental morphologies observed in the Hainin fauna suggests different interrelationships than previously suggested. In particular, the genus Proremiculus is considered morphologically intermediate between Afrodon and Remiculus, and the latter is no longer recognised as the sister group of Adapisoriculus. Although the highest diversity of adapisoriculids occurs in Europe, the oldest and most primitive members of the family were found in India and Africa, respectively. The geographic origin of the family could thus be located in any of these three continents, depending on the importance attributed to each of these factors. The coexistence of primitive and derived adapisoriculids at Hainin might indicate a very quick diversification in Europe, probably starting around the K−T boundary.
The phylogenetic relationships of the Paleocene crocodylian Akanthosuchus langstoni are assessed using published data matrices and morphological data from the holotype and referred specimens. Cladistic analyses indicate that Akanthosuchus is unequivocally nested within Alligatoroidea. Weak support from a majority rule consensus tree indicates that Akanthosuchus may be more closely allied with alligatorines than with caimanines, but in the strict consensus tree these relationships remain ambiguous. There is no evidence from phylogenetic analyses to support the hypothesis that Akanthosuchus represents the postcrania of the Paleocene crocodylians Navajosuchus or Ceratosuchus. Growth marks observed in histological sections of osteoderms of the holotype of Akanthosuchus langstoni indicate that it was at least eight years old at the time of death. Although the individual may not have been fully mature at the time of death, lineage dwarfism cannot be ruled out as a possible reason for its relatively small size.
Originally designated by Dollo in 1907, the holotype of Eosuchus lerichei has never been carefully described but simply cited and compared in a number of papers. This work is an attempt to fill this gap and to place this taxon in a cladistic phylogenetic context. E. lerichei can be considered a valid basal gavialoid from late Paleocene of North Western Europe, sharing the presence of extremely enlarged foramina aerea on quadrates with the coeval Eosuchus minor from eastern North America (formerly described as Gavialis minor). These two species can be considered sister taxa and, for priority reason, they should be both ascribed to genus Eosuchus. The results of the cladistic analysis show that the European species possess characters that can be considered as slightly derived if compared to those of its American relative, suggesting an eastward dispersion from North America before the Paleocene–Eocene boundary and before the full opening of the Atlantic Ocean or local evolution from a basal gavialoid stock similar to E. minor. Both species of Eosuchus come from marine outcrops and represent a further evidence for the salt−water tolerance of the earliest stages of Gavialoidea evolutionary history. Despite the present endemicity of the only living gharial, Gavialis gangeticus, the historical biogeography of gavialoids shows a lost global distribution and reveals several transoceanic dispersals.
A new genus and species belonging to Dyrosauridae, Arambourgisuchus khouribgaensis, from the Thanetian (Palaeocene) of Morocco, is erected. Two more or less complete skulls and three mandibular fragments enable a reconstruction of the anatomical characteristics of this species. Dyrosaurid systematics is mainly based on mandibular characters. The comparison of this new material with several dyrosaurid species previously known provides new systematic data for this group. The width of the interfenestral bar, the shape and development of the occipital tuberosities and the shape of the supraoccipital and the basioccipital are of particular importance. A phylogenetic analysis of the dyrosaurids provides an outline of the relationships between the best known species. Chenanisuchus lateroculi is the most primitive dyrosaurid. Sokotosuchus ianwilsoni and Phosphatosaurus gavialoidesform a clade, more closely related to other dyrosaurids than to Chenanisuchus lateroculi. The relationships between Arambourgisuchus, Rhabdognathus, Congosaurus, and Hyposaurus are unclear, and the two latter taxa remain too poorly known to provide an uncontested phylogenetic result. The dyrosaurids are known from nearly all continents. The phylogenetic results suggest a North African range for basal members, and the wide distribution of Rhabdognathus and Hyposaurus confirms the possibility of transoceanic dispersal of these taxa. Unfortunately, many dyrosaurids are insufficiently known to be included in the analysis, and the present analysis considers mainly African forms. A better knowledge and the inclusion of other taxa from other geographic regions should significantly improve and modify the hypothesis.
A new plesiadapiform primate, Phoxomylus puncticuspis gen. et sp. nov., is described based on an isolated but well−preserved upper molar from the early Tiffanian (late Paleocene) Cochrane 2 locality, southwestern Alberta, Canada. Although possessing a robust postprotoconal fold, an unambiguous synapomorphy of primates, Phoxomylus differs from other plesiadapiforms in its retention of primitive molar features, including acutely pointed major cusps and sharp crests, deep trigon basin, and lack of the bunodont coronal specializations that purportedly marked the transition from insectivorous non−primate ancestors to omnivorous/frugivorous basal primates. Coronal features of the holotype of P. puncticuspis imply that during mastication the mandible was adducted in a near−vertical plane, with little capacity for the transverse movement that is already seen in molar morphology of the earliest and most basal plesiadapiform, Purgatorius. Instead, molar morphology in P. puncticuspis implies emphasis on vertical piercing and shearing, specializations for insectivory unlikely to have been derived via reversal from plesiadapiform ancestors having more bunodont molars adapted for omnivory/frugivory. If that is the case, a long “ghost lineage” must link P. puncticuspis to other, basal plesiadapiforms that have yet to be discovered but that had not yet evolved omnivorous adaptations of the dentition.
A second species of the microchoerine omomyid genus Melaneremia, M. schrevei sp. nov. is described. It has been collected from the upper shelly clay unit of the Woolwich Formation, earliest Ypresian, Eocene, of Croydon, Greater London, UK. Phylogenetic analysis shows M. schrevei to be the most primitive member of the main clade of the Microchoerinae and demonstrates the initial dental evolution that separated this European subfamily from other omomyids. Calibration of the Woolwich upper shelly clay unit to the later part of the Paleocene–Eocene Thermal Maximum shows that speciation leading to the Microchoerinae took place within 170 ky of the beginning of the Eocene. Tentative identification of M. schrevei in the Conglomérat de Meudon of the Paris Basin suggests close time correlation with the upper part of the Woolwich Formation.
Vastan Lignite Mine in southeastern Gujarat, India, produces the oldest known Cenozoic land−mammals and the only early Eocene continental vertebrate fauna known from India (e.g., Bajpai et al. 2005; Rana et al. 2005, 2008; Rose et al. 2006, 2008, 2009; Smith et al. 2007; Rage et al. 2008). The fauna comes from the Cambay Shale Formation and has been dated as middle Ypresian (~52 Ma, early Cuisian) based on a common nummulitid foraminiferan from about 15 m above the vertebrate−producing layer (Sahni et al. 2006; Rana et al. 2008). However, a recent study of dinoflagellate cysts from the section suggests that the deposits may be as old as 54–55 Ma (Garg et al. 2008). Although some elements of the fauna, such as anthracobunids and lagomorphs, have Asian affinities, a surprising number of taxa among the snakes, bats, insectivores, primates, rodents, and artiodactyls appear to be most closely related to early Eocene European or North American taxa. This may simply reflect the poor state of knowledge of contemporary south Asian vertebrate faunas; alternatively, it might be evidence of previously unsuspected early Eocene faunal exchange between Europe and southwest Asia. We report here two teeth of a tillodont from Vastan Mine, which constitute the first record of the mammalian order Tillodontia known from India. Despite the much greater generic diversity of tillodonts in Asia than elsewhere, the Vastan tillodont shows clear affinities with Euramerican esthonychines.
Isolated cheek teeth from the late Puercan (early Paleocene) Split Lip Flats local fauna, from the head of Willow Wash, Nacimiento Formation, San Juan Basin, represent a new genus and species of hyopsodontid “condylarth”, Chacomylus sladei. The teeth are small, bunodont, and are remarkable for the enlargement of the protocone, pronounced exodaenodonty, and high degree of attritional wear caused by transverse shearing or grinding. Although C. sladei bears a superficial resemblance to the apheliscine apheliscid Gingerichia, a preliminary phylogenetic analysis confirms that it is a hyopsodontid, closely related to the Puercan hyopsodontids Valenia wilsoni and Litomylus orthronepius.
We describe an additional fragmentary upper molar and the first lower molar known of Monotrematum sudamericanum, the oldest Cenozoic (Paleocene) monotreme. Comparisons suggest that the monotreme evolution passed through a stage in which their molars were “pseudo−triangulate”, without a true trigonid, and that the monotreme pseudo−triangulate pattern did not arise through rotation of the primary molar cusps. Monotreme lower molars lack a talonid, and consequently there is no basin with facets produced by the wearing action of a “protocone”; a cristid obliqua connecting the “talonid“ to the “trigonid” is also absent. We hypothesize that acquisition of the molar pattern seen in Steropodon galmani (Early Cretaceous, Albian) followed a process similar to that already postulated for docodonts (Docodon in Laurasia, Reigitherium in the South American sector of Gondwana) and, probably, in the gondwanathere Ferugliotherium.
New dental and postcranial remains of the alleged louisinine hyopsodontid “condylarth” Microhyus from the European Paleocene/Eocene transition are described, and prompt a reevaluation of the genus. New specimens belonging to Microhyus musculus from Dormaal (MP7, Belgium) provide the first evidence of the lower dentition of the type species. We describe M. musculus? from Pourcy (MP7, France) and cf. Microhyussp. from Berru (MP6a, France). A rich original assemblage of M. reisi from Silveirinha (MP7, Portugal) allows a detailed description of the morphological dental variation within that species. Well−preserved astragali and calcanei from Silveirinha can be confidently attributed to Microhyus reisi. Functional analysis of these elements suggests that Microhyus was a terrestrial mammal capable of rapid running or jumping. The pedal morphology of Microhyus is very similar to that of Paschatherium. These louisinines share some derived characters with the hyopsodontids Apheliscus and Haplomylus (e.g., the occurrence of a cotylar fossa on the astragalus) but they differ from Hyopsodus. Therefore, in view of the pedal morphology alone, the hyopsodontids may be polyphyletic. Given the dental similarities between Microhyus and the early representatives of the order Macroscelidea, we compared the tarsal morphology of louisinines with that of modern macroscelidids (Paleogene tarsal remains are currently unknown for this group). Macroscelidids and louisinines present some similarities in their astragalar morphology; however, the macroscelidid astragalus appears to be too specialized to be compared with that of Microhyus and Paschatherium.
Nyctitheriids are primitive insectivores that were relatively abundant and diverse in North America and Europe during the middle Paleocene through to the middle Oligocene. The nyctitheriids from Asia are poorly known and show several distinctive characters. Here we describe the late Paleocene Asionyctia guoi gen. et sp. nov., the first fairly well known Asian nyctitheriid, from the Subeng locality near the city of Erlianhot (Erenhot) in Inner Mongolia, China. Among its most conspicuous features are the paraconid positioned high on p4, the rather primitive morphology and size of p3, the premolariform P4/p4 and the transverse upper molars with a small, straight postcingulum. Except for the paraconid positioned high on p4, these combined features are also present in other Asian nyctitheriids, but absent in North American or European forms. We performed a cladistic analysis, based on a set of 20 dental characters, to resolve higher−level phylogenetic relations within Nyctitheriidae. The strict consensus tree groups all Asian forms in a single clade, for which we propose the rank of a subfamily and the name Asionyctiinae subfam. nov. Within Nyctitheriidae, a semimolariform P4/p4, as in Leptacodon tener, is considered primitive, and we consider the morphologically simplified P4/p4 of Asionyctiinae derived within Nyctitheriidae. Asionyctiinae can be derived from an American, primitive Leptacodon−like ancestor migrating into Asia, with the reduction of P4/p4 occurring on the Asian continent. Considering the derived morphology and the relatively high diversity of Asionyctiinae during the Asian late Paleocene, and the inferred conservative nature of the family Nyctitheriidae, we suggest an early Tiffanian time for the migration of nyctitheriids into Asia.
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