New Alicornops (Rhinocerotidae) remains from Lower and Middle Siwaliks,Pakistan

Unpublished rhinocerotid remains from the Lower and Middle Siwaliks of Pakistan are described in this paper and recognized as two species of Alicornops. Alicornops complanatum (Heissig) is identified in the Dhok Pathan Formation of the Middle Siwaliks and Alicornops laogouense Deng in the Kamlial Formation of the Lower Siwaliks. The Dhok Pathan Formation levels with A. complanatum are roughly correlated with the late Miocene-Pliocene European mammal zones MN10-15. In turn, levels with A. laogouense of the Kamlial Formation would correlate with the middle–late Aragonian (middle Miocene) European MN5. The recognition of the Chinese species A. laogouense in the Potwar Plateau represents the first discovery of this taxon in Pakistan and increases the geographical and stratigraphic distributions of this species, and adds to the rhinocerotid association from the Siwaliks. In turn, the presence of A. complanatum in the Siwaliks of Potwar Plateau also enlarges its geographic distribution in Pakistan, as it was previously known from the Bugti Hills of Balouchistan. The absence of Alicornops from the Siwaliks in the Chinji and Nagri formations (between late MN5 and MN9 zones) might be due to an inadequate fossil record, as other rhinocerotid species are known from Kamlial to Dhok Pathan formations. However, the two recorded species of Alicornops could also reveal two independent migration waves as supported by the appearance of other taxa in different formations. A summary of fossil Cenozoic rhinocerotids from different areas of Pakistan is also presented.     

A new species of Golunda (Rodentia, Muridae) from the Late Pleistocene of Indian Himalaya

The youngest fossil Golunda (Rodentia, Muridae) is described from the Late Pleistocene fluvio-lacustrine deposits, exposed at Dulam (Bageshwar), Kumaun Lesser Himalaya, India. The age of fossiliferrous horizon is estimated as 31,000 yr BP. A new species, Golunda dulamensis nov. sp. has highly derived characters, e.g., antero-posteriorly stretched molars, upper molars with more length and less width, stephanodonty, cusps in M₃ strongly inclined backward giving the molars a very stretched aspect, and metaconid and entoconid in M₃ forming almost straight lingual row of the cusps. G. dulamensis nov. sp. is most similar to present day G. ellioti but differs from the later slightly by larger size, a thin connection between t₄ and t₅ in M¹, and comparatively larger entoconid and very weakly developed antero-labial cusp in M₃. We suggest that highly specialized molars of G. dulamensis nov. sp. and present day G. ellioti are derivable through G. kelleri. We also propose that Golunda migrated from Asia to Africa, not from Africa to Asia as was thought by earlier workers.     

Early Paleogene decapod crustaceans from the Sulaiman and Kirthar Ranges,Pakistan

A set of Paleocene and Eocene decapod crustaceans is described from the Sulaiman and Kirthar Ranges of Pakistan. The fossil crabs Proxicarpilius planifrons Collins and Morris, 1978 and Pakicarcinus orientalis (Collins and Morris, 1978), already known in the Eocene of northern Pakistan, are reported for the first time in the Paleocene of southern Pakistan, enlarging the stratigraphic and the palaeobiogeographical ranges of these species. The callianassid genus Calliax de Saint Laurent, 1973 is reported for the first time in the Paleocene of southern Pakistan; this is the oldest record for the genus.     

Diversity of Menispermaceae from the Paleocene and Eocene of South China

We present here the earliest known Asian fossil records of the Menispermaceae based on fossil fruits from Paleocene and Eocene localities in South China. A new genus and species, Paleoorbicarpum parvum sp. nov., and two new species of Stephania Loureiro, S. ornamenta sp. nov. and S. geniculata sp. nov., are recognized from Paleocene deposits of the Sanshui Basin, Guangdong, and a new occurrence of the widespread Eocene species Stephania auriformis (Hollick) Han & Manchester is recognized from the Maoming Basin, Guangdong. The Paleocene Stephania specimens described here represent the earliest fossil endocarp record of the Menispermaceae in eastern Asia. This discovery shows that the moonseed family had arrived in tropical and humid South China by at least the middle Paleocene, which provides important evidence for the origin and phytogeographic history of the family.     

A new basal testudinoid turtle (Testudinoidea: “Lindholmemydidae”) from the Upper Paleocene of Mongolia

A new genus and species of a basal testudinoid turtle of the family “Lindholmemydidae,” Paramongolemys khosatzkyi gen. et sp. nov., is described based on shell material from the Naranbulak Formation (Upper Paleocene) of the Naran Bulak and Khaychin-Ula IE localities (Mongolia). Material of this species includes some specimens previously described as Mongolemys tatarinovi.     

The fossil Bovidae (Artiodactyla, Mammalia) from Gesher Benot Ya'aqov, Israel: Out of Africa during the Early-Middle Pleistocene transition

We report the study of the collection of fossil bovid specimens from the Early-Middle Pleistocene Acheulian site of Gesher Benot Ya‘aqov. This locality, situated in the Levantine Corridor (the bottleneck that connects Africa and Eurasia) is a key site to explain the faunal and human dispersals out of Africa during the Matuyama/Brunhes boundary around 0.8 Ma. Two species of bovine (Bos sp., and Bovini gen. et sp. indet. cf. Bison sp.), one antelope (Gazella sp. cf. G. Gazella), and another indeterminate Bovidae gen. et sp. indet., have been recorded. The largest species, Bos sp., is an African immigrant related to the species from the Eritrean site of Buia, Bos buiaensis, which evolved from the buffalo of Olduvai Pelorovis oldowayensis, and colonized the Eurasian continent in parallel with the dispersal of the Acheulian culture into the northern continent. Numerous important species first recorded in several localities of Early-Middle Pleistocene transition from Eurasia are included in this dispersal out of Africa, including the megaherbivore, Palaeoloxodon antiquus, and the carnivores Crocuta crocuta, and later, Panthera leo and Panthera pardus. This faunal turnover is coincident with the change to colder climates that dominated the Middle Pleistocene.     

Pleuromeia discovered from the Middle Triassic Linjia Formation of Benxi,Northeast China

A new species Pleuromeia shaolinii Zhang et Wang n. sp. was established under the family Pleuromeiaceae, based on eight specimens of stems and leaves, cones, rhizophores, and adventitious roots that were recently collected from the Middle Triassic Linjia Formation in Benxi, Northeast China. P. shaolinii n. sp. is similar to the type species P. sternbergii, in the morphology of the stem and rhizophore, but differs from it by the transversely rounded sporophylls organized in a heterosporous cone, and the two conspicuous triangular scars present on both sides of the vascular bundle scar. The Linjia flora, an only reliable Middle Triassic flora in Northeast China, was discovered 40 years ago. Although the Linjia Formation can be compared with the Middle Triassic Ermaying Formation from another area of the North China plate, its age was questionable due to the absence of the Early to Middle Triassic index fossil Pleuromeia. Based on the flora similarity, the zircon dating, lithologic characteristics, presence of a disconformity, and the occurrence of the Middle Triassic–Early Cretaceous insect fossil Sogdoblatta, the age of the Linjia Formation can be confirmed as Middle Triassic. The new species not only supplements our knowledge in biology and taxonomy of Pleuromeia, but also supports the Middle Triassic age assignment of the Linjia flora.     

The first species of Hapalodectes (Mesonychia,Mammalia) from the middle Paleocene of China (Qianshan Basin,Anhui Province) sheds light on the initial radiation of hapalodectids

A lower jaw of the mesonychian Hapalodectes is reported from Nongshanian sediments (Upper Doumu Formation; middle Paleocene) of the Qianshan Basin (Anhui Province, China). The fragmentary mandible is only the third specimen of Hapalodectidae discovered in Paleocene deposits, and the first in south east China; it is moreover the oldest, the two other specimens having been found in Gashatan (late Paleocene) localities. The premolars and molars of the new fossil are morphologically similar to Hapalodectes dux (late Paleocene of Mongolia), which has been considered to be the most primitive hapalodectid, but their relative proportions recall H. paleocenus and the Eocene Hapalodectes species. As a result, the fossil described herein appears to be different from the other previously described species of Hapalodectes in being morphologically intermediate between H. dux and the other Hapalodectes species, notably the Bumbanian Hapalodectes hetangensis and H. huanghaiensis from China; it is thus identified as a new species, Hapalodectes lopatini (possibly a male individual). Its discovery is important because it sheds light on the initial radiation of hapalodectids. The presence of one primitive hapalodectid in Mongolia previously suggested the Mongolian Plateau as the centre of origination of this carnivorous family, but the discovery of H. lopatini in older sediments from south‐east China challenges this hypothesis. In the earliest Eocene, Hapalodectes dispersed from Asia to North America; this event being part of the ‘East of Eden’ dispersals. This event resulted in the geographical separation of two distinct Hapalodectes groups, in North America and south‐eastern China respectively.     

Origin of the white shark Carcharodon (Lamniformes: Lamnidae) based on recalibration of the Upper Neogene Pisco Formation of Peru

Abstract: The taxonomic origin of the white shark, Carcharodon, is a highly debated subject. New fossil evidence presented in this study suggests that the genus is derived from the broad‐toothed ‘mako’, Carcharodon (Cosmopolitodus) hastalis, and includes the new species C. hubbelli sp. nov. – a taxon that demonstrates a transition between C. hastalis and Carcharodon carcharias. Specimens from the Pisco Formation clearly demonstrate an evolutionary mosaic of characters of both recent C. carcharias and fossil C. hastalis. Characters diagnostic to C. carcharias include the presence tooth serrations and a symmetrical first upper anterior tooth that is the largest in the tooth row, while those indicative of C. hastalis include a mesially slanted third anterior (intermediate) tooth. We also provide a recalibration of critical fossil horizons within the Pisco Formation, Peru using zircon U‐Pb dating and strontium‐ratio isotopic analysis. The recalibration of the absolute dates suggests that Carcharodon hubbelli sp. nov. is Late Miocene (6–8 Ma) in age. This research revises and elucidates lamnid shark evolution based on the calibration of the Neogene Pisco Formation.     

Bridging the extant and fossil record of planktonic foraminifera: implications for the Globigerina lineage

We conducted a morphometric study and wall texture analysis on extant and fossil specimens of the planktonic foraminifera Globigerina falconensis plexus. Our global data reveal morphological inconsistencies between fossil and extant populations. Our results are significant as G. falconensis is widely used in palaeoceanographic studies in conjunction with its sister taxon G. bulloides. Morphologically these two species are similar, with the main difference being the distinctive apertural lip present in G. falconensis. We selected cores covering the entire stratigraphic range of G. falconensis, from the early Miocene to current day, spanning sites from high latitudes in the North Atlantic Ocean and the southern Indian Ocean to sites in equatorial regions. The morphology found in the modern ocean is not consistent with the Miocene holotype of Globigerina falconensis Blow described from lower Miocene sediments in Venezuela. A more lobate morphology evolved in the late Miocene, thus, a new name is required for this morphotype, coexisting in the modern oceans with G. falconensis s.s. We thus describe the new morphospecies, G. neofalconensis for the more lobate forms which evolved in the late Miocene and inhabit the modern oceans. Additionally, we report a pseudocancellate wall texture present in the G. falconensis plexus. We use the molecular sequences from the PR² database to explore the generic attribution of the G. falconensis lineage, confirming its close relationship with G. bulloides and its retention in the genus Globigerina.     

Exploring the Formation of a Disjunctive Pattern between Eastern Asia and North America Based on Fossil Evidence from Thuja (Cupressaceae)

Thuja, a genus of Cupressaceae comprising five extant species, presently occurs in both East Asia (3 species) and North America (2 species) and has a long fossil record from Paleocene to Pleistocene in the Northern Hemisphere. Two distinct hypotheses have been proposed to account for the origin and present distribution of this genus. Here we recognize and describe T. sutchuenensis Franch., a new fossil Thuja from the late Pliocene sediments of Zhangcun, Shanxi, North China, based on detailed comparisons with all living species and other fossil ones, integrate the global fossil records of this genus plotted in a set of paleomaps from different time intervals, which show that Thuja probably first appeared at high latitudes of North America in or before the Paleocene. This genus reached Greenland in the Paleocene, then arrived in eastern Asia in the Miocene via the land connection between East Asia and western North America. In the late Pliocene, it migrated into the interior of China. With the Quaternary cooling and drying, Thuja gradually retreated southwards to form today’s disjunctive distribution between East Asia and North America.     

Les foraminifères du Paléocène et de l'Éocène basal du sillon nord-pyrénéen (Aquitaine, France)

This paper presents for the first time the inventory of the Paleocene and Lower Eocene foraminifers located in the North Pyrenean trough, between the Atlantic Ocean and the neighbourhood of the town Pau. They have been studied from three outcrops. The Bidart Beach section shows the Lasseube Formation from the Cretaceous/Paleogene boundary to the base of the P 3a zone. The Loubieng Quarry section, near Orthez, represents the upper P 3a zone and the lower P 3b zone as well as the Lasseube Formation / Pont Labau Formation boundary. The interval between the upper part of the P 3b zone and the upperest part of the P 5 zone crops out along the Gan - Rébénacq road with a hiatus located at the Paleocene / Eocene boundary, the whole interval belonging to the Pont Labau Formation. 394 taxons of foraminifers are present in this formations: 349 benthic and 45 planktonic species. The Velasco type benthic foraminifers show a middle bathyal depositional environment, with a paleobathymetry included between 500-600 m and 1000 m: Nuttallides truempyi, Osangularia velascoensis, Bulimina trinitatensis. The Midway type species which were transported by the turbidite currents from the lower to middle neritic environments are frequent as well as the Cretaceous reworked species. The species number is low: 29 in the iridium layer of the Cretaceous/Paleogene boundary (P 0 zone). Fauna grows rich quickly in the Pα zone reaching 129 species. The diversity grows up progressively from P α to the P 4a zone (NP 8). The disappearances are rare until his horizon, but their number is bigger than the number of appearances from the P 4b zone. It reaches their maximum between the P 4c zone and the P 5 zone. It shows that the decline of the Paleocene fauna begins around 2 million years before the thermal event of the Paleocene / Eocene boundary in the Aquitaine sections. The disappearances stay important in the Lower Eocene - Ypresian, but the appearance of lots of Eocene species show that the fauna renewal is located in this stage. The Cretaceous taxa dominate in the Paleocene benthonic fauna. The appearance or the disappearance of some species has a stratigraphic value in the Aquitaine region: the disappearance of Coryphostoma incrassata in the P 1b zone, the appearances of Plectina dalmatina, Elongobula grata (P α); Bulimina tuxpamensis (P 2); Tritaxilina cubensis, Thalmannita madrugaensis (P 3a); Svenia bulbosa (P 3b); Discorbis perovalis (P 4a/NP 7); Elongobula pulchra, E. pupa, Asterigerina bartoniana, Neorotalia gr. tuberculata (P 4a/NP 8); Bigenerina pannonica, Pentellina pseudosaxorum (P 5/NP 9-10).     

New skull of Lesothosaurus (Dinosauria: Ornithischia) from the Upper Elliot Formation (Lower Jurassic) of southern Africa

A new ornithischian skull from the Elliot Formation of southern Africa is described. The specimen is compared in detail with the fabrosaurid Lesothosaurus diagnosticus. It actually shares many characters with specimens of the syntypes of this species or specimens referred to it. It is nevertheless not identical to any of these specimens and it is, moreover, remarkably larger than them. The possibility of attributing this specimen to a so far undescribed ‘large fabrosaur’ from the same formation is discussed. It is concluded that the specimen in question in this paper, while being ascribable to the genus Lesothosaurus, cannot be determined to a specific level until the existence of two fabrosaurid species in the ‘Stormberg Group’ is demonstrated and their range of morphological and size variation is properly appraised.     

The oldest Odontostomidae (Mollusca: Gastropoda): Bahiensis priscus n.?sp. (Paleocene, Uruguay)

The land snail family Odontostomidae has a poor fossil record, mainly from the middle Paleogene and early Neogene of Argentina. In this paper a new species of Odontostomidae from the Paleocene of Uruguay (Queguay Formation) is described. Bahiensis priscus n.?sp. represents the first record of the genus Bahiensis Jousseaume 1877, and the oldest record for an Odontostomidae. The new species is characterized by a pupoid fusiform shell and an oval aperture with a single axial columellar fold. Present distribution of the genus indicates a tropical?Csubtropical environment, in high humidity rate areas.     

?Kenyaichthyidae fam. nov. and ?Kenyaichthys gen. nov. – First Record of a Fossil Aplocheiloid Killifish (Teleostei,Cyprinodontiformes)

The extant Cyprinodontiformes (killifishes) with their two suborders Cyprinodontoidei and Aplocheiloidei represent a diverse and well-studied group of fishes. However, their fossil record is comparatively sparse and has so far yielded members of the Cyprinodontoidei only. Here we report on cyprinodontiform fossils from the upper Miocene Lukeino Formation in the Tugen Hills of the Central Rift Valley of Kenya, which represent the first fossil record of an aplocheiloid killifish. A total of 169 specimens - mostly extraordinarily well preserved - and a sample of ten extant cyprinodontiform species were studied on the basis of morphometrics, meristics and osteology. A phylogenetic analysis using PAUP was also conducted for the fossils. Both the osteological data and the phylogenetic analysis provide strong evidence for the assignment of the fossils to the Aplocheiloidei, and justify the definition of the new family †Kenyaichthyidae, the new genus †Kenyaichthys and the new species †K. kipkechi sp. nov. The phylogenetic analysis unexpectedly places †Kenyaichthys gen. nov. in a sister relationship to the Rivulidae (a purely Neotropical group), a probable explanation might be lack of available synapomorphies for the Rivulidae, Nothobranchiidae and Aplocheilidae. The specimens of †K. kipkechi sp. nov. show several polymorphic characters and large overlap in meristic traits, which justifies their interpretation as a species flock in statu nascendi. Patterns of variation in neural and haemal spine dimensions in the caudal vertebrae of †Kenyaichthys gen. nov. and the extant species studied indicate that some previously suggested synapomorphies of the Cyprinodontoidei and Aplocheiloidei need to be revised.     

Taxonomy and paleobiogeographic implication of Glabrobournonia Morris and Skelton (Hippuritida,Radiolitidae) from the Late Cretaceous Yigeziya Formation,southwestern Tarim Basin

A re-examination of the specimens that were identified as Biradiolites minor Pojarkova from the late Campanian to early Maastrichtian middle member of the Yigeziya Formation of southwestern Tarim Basin reveals that they should be assigned to the genus Glabrobournonia Morris and Skelton. Glabrobournonia is a genus of radiolitids characterized by indented radial bands, salient ridges on the shell margins and absence of fine ribs on the surface of the right valve. Apart from southwestern Tarim Basin, Glabrobournonia minor (Pojarkova) has also been recorded from the late Campanian of Fergana and Alai basins. The central Asian, late Campanian to early Maastrichtian G. minor differs from the late Campanian to Maastrichtian, eastern Arabian type species Glabrobournonia arabica Morris and Skelton in the flat left valve and an additional fourth ridge on the junction of the dorsal and posterior sides of the right valve. Biradiolites ingens (Des Moulins) could be the direct ancestor of Glabrobournonia. The paleogeographic distribution of Glabrobournonia suggests that this genus dispersed to central Asia from the late Campanian time, becoming widely distributed in the eastern Tethyan region rather than endemic to eastern Arabia. Correspondingly, specimens belonging to Gyropleura yielded from the same bed as G. minor in southwestern Tarim Basin, are similar to the specimens which were attributed to the eastern Arabian Gyropleura sp.; Campanian to early Maastrichtian Osculigera specimens described from the Yigeziya Formation are comparable with those known from the Campanian–Maastrichtian of Iran, Afghanistan and eastern Arabia. The similarity of the rudist assemblages between central Asia and eastern Arabia suggests a faunal connection and affinity between the north and south margins of the eastern Tethyan realm during Campanian to early Maastrichtian times.     

Anthracotheriid artiodactyl Anthracokeryx and an upper Eocene age for the Youganwo Formation of southern China

Abstract

The two recently collected mammalian dentary fragments from the Eocene Youganwo Formation of Guangdong Province, southern China, are referred to the anthracotheriid species Anthracokeryx naduongensis based on phylogenetic analysis and size comparison. One of these specimens (SYSU-M-1) is the first mammal fossil described from the Youganwo Formation. It was attributed previously to the perissodactyl genus Lunania. Anthracokeryx naduongensis was described originally from the lower upper Eocene Na Duong Formation in Northern Vietnam. The second record of this species supports a basal upper Eocene correlation for the Youganwo Formation, which was estimated previously as middle or late Eocene.     

Early Miocene cricetids (Rodentia) from the Junggar basin (Xinjiang, China) and their biochronological implications

Among the 14 small mammal species from the early Miocene locality of the northern area of the Junggar basin (northern Xinjiang, China), four species are cricetids (Rodentia): unnamed species of Cricetodon and Eumyarion, and two new species, Karydomys debruijni nov. sp. and Megacricetodon beijiangensis nov. sp. Some aspects of the morphology of Cricetodon sp. are shared by Eucricetodon from the late Oligocene, suggesting that these specimens could be of intermediate form between Eucricetodon and Cricetodon. One tooth of Eumyarion sp. was found, making its determination uncertain, but its morphology is clearly differentiated from the one of Cricetodon sp. The species K. debruijni nov. sp. is established based on its primitive features compared to the species known in Europe and Anatolia, and its specific association of characters compared to Karydomys dzerzhinskii. M. beijiangensis nov. sp. shows many plesiomorphic features compared to the species already described in the middle Miocene of China. Based on both the whole assemblage of rodents and the species of cricetids, the biochronologic position and the age of the locality are discussed. The locality appears to be biochronologically very close to the fauna from the Chul’adyr Formation in Aktau Mountains, but we propose an age slightly older than the one proposed for this fauna, probably equivalent to the MN3 biozone in Europe.