A new species of the genus Microbunodon (Anthracotheriidae, Artiodactyla) from the Miocene of Pakistan: genus revision, phylogenetic relationships and palaeobiogeography

Lihoreau, F., Blondel, C., Barry, J. & Brunet, M. (2004). A new species of the genus Microbunodon (Anthracotheriidae, Artiodactyla) from the Miocene of Pakistan: genus revision, phylogenetic relationships and palaeobiogeography. — Zoologica Scripta , 33 , 97–115.
New unpublished remains of small Anthracotheriinae are described. First, materials from the upper Oligocene (MP 30) locality of La Milloque, southwest France, permit a review of the species Microbunodon minimum . Thereafter, fossils from the middle and late Miocene of the Potwar Plateau, Pakistan are attributed to the European genus Microbunodon . Microbunodon milaensis sp. n. from the Nagri Formation (between 10.3 and 9.2 Ma), Pakistan, is described and the species M. silistrensis from the Lower Manchar Formation (between 16 and 15 Ma) and from the Chinji Formation (between 12.7 and 11.5 Ma), Pakistan, is reviewed. The new species represents the last occurrence of the subfamily Anthracotheriinae, around 9.3 Ma. Similar materials from the Bugti and Siwalik Hills were previously considered as a small Anthracotherium . Comparisons with M. minimum from the European late Oligocene lead to a complete revision of the genus and permit definition of a new set of characters, which separate Microbunodon from Anthracotherium . A cladistic analysis reconsiders phylogenetic relationships among Anthracotheriinae, separating an Anthracothema–Anthracotherium clade and an Anthracokeryx–Microbunodon clade. Microbunodon appears to stem from the Asian late Eocene–lower Oligocene genus Anthracokeryx . These results imply a new distribution of the genus Microbunodon showing exchanges between Europe and Asia during the late Oligocene and probably the lower Miocene.     

Sexual and individual morphometric variation in Libycosaurus (Mammalia, Anthracotheriidae) from the Maghreb and Libya

Anthracotheres of late Middle Miocene and Late Miocene age have been described from several localities in northern Africa, all of them currently assigned to the genus Libycosaurus Bonarelli, although in several previous works they were assigned to Merycopotamus Falconer and Cautley, a considerably younger and specialised form from the Indian subcontinent which has quite different dental and cranial morphology. Three species of Libycosaurus have been named, but there has been some doubt about the morphometric variation within the various species, with some authors such as Gaziry (1987) placing the fossils from Sahabi (Latest Miocene, Libya) and Beglia (end Middle Miocene to basal Late Miocene, Tunisia) into the same species despite marked size differences, and others (Ducrocq et al., 2001) creating a species for a restricted sample of small specimens from Nementcha (Late Middle Miocene, Algeria), but which overlaps with the range of size variation of Beglia fossils. The aim of this paper is to examine the available samples in greater depth in order to understand the morphometric variation in these anthracotheres. It is confirmed that the Beglia sample is quite variable (Black, 1972), both morphologically and metrically, but it is concluded that it nevertheless belongs to a single species, because specimens from some of the localities within the Beglia Formation (e.g. Loc. 17 in the lower levels at Beglia) span the entire range of variation (Pickford, 1994). The sample from Nementcha cannot be distinguished from the Beglia sample on any consistent metric or morphological basis, but in general the specimens fall at the low end of the range of variation of the Beglia sample. It is thus likely that L. algeriensis is a synonym of L. anisae. The Sahabi and Chad samples (L. petrocchii), in contrast, fall above the known range of variation of the Beglia material in almost all metric features, but are close to it morphologically, and they are considered to represent a species distinct from the Beglia sample.     

New observations on the Late Miocene–Early Pliocene Lutrinae (Mustelidae: Carnivora,Mammalia) from the Middle Awash,Afar Rift,Ethiopia

New observations on the Late Miocene and Earliest Pliocene mustelids from the Middle Awash of Ethiopia are presented. The Middle Awash study area samples the last six million years of African vertebrate evolutionary history. Its Latest Miocene (Asa Koma Member of the Adu-Asa Formation, 5.54–5.77 Ma) and Earliest Pliocene (Kuseralee and Gawto Members of the Sagantole Formation, 5.2 and 4.85 Ma, respectively) deposits sample a number of large and small carnivore taxa among which mustelids are numerically abundant. Among the known Late Miocene and Early Pliocene mustelid genera, the Middle Awash Late Miocene documents the earliest Mellivora in eastern Africa and its likely first appearance in Africa, a new species of Plesiogulo, and a species of Vishnuonyx. The latter possibly represents the last appearance of this genus in Africa. Torolutra ougandensis is known from both the Late Miocene and Early Pliocene deposits of the Middle Awash. The genus Sivaonyx is represented by at least two species: S. ekecaman and S. aff. S. soriae. Most of the lutrine genera documented in the Middle Awash Late Miocene/Early Pliocene are also documented in contemporaneous sites of eastern Africa. The new observations presented here show that mustelids were more diverse in the Middle Awash Late Miocene and Early Pliocene than previously documented.     

内蒙古宝格达乌拉晚中新世副竹鼠化石(英文)

描述了1996-2007年间在内蒙古阿巴嘎旗晚中新世宝格达乌拉组采集的副竹鼠化石。材料包括下颌骨1件、牙齿14枚和肢骨4件,分别采自该组的2个地点。根据标本较大的尺寸、形态特征,以及颊齿的釉质曲线高度,宝格达乌拉材料被归入三趾马层副竹鼠Pararhizomys hipparionum Teilhard de Chardin Young,1931。Pararhizomys是一类具高冠、脊形和冠面构造简单颊齿的啮齿动物。该属目前发现的化石地点少,材料也不多,主要集中于中国北方和蒙古高原周缘,其地史分布为晚中新世早期至上新世中期。与鼢鼠类(siphneids)和鼠平类(arvicolids)一样,Pararhizomys属的颊齿有由低冠到高冠发展的趋势,具体表现为牙齿侧面釉质曲线(dentine tract)由平直、远离内外两侧谷底到起伏、接近甚至超过内外两侧谷底。为方便比较,本文为釉质曲线的高度设立"H"指数,即从每侧最深谷的谷底和釉质曲线的最高处取平行于冠面的平行线,此两平行线之间的垂直距离为"H"。H值越大,表明齿冠越低,反之则齿冠越高。尽管目前发现的副竹鼠标本不多,但似乎可以看到从早期到晚期,其颊齿的个体有从小变大,釉质曲线高度H值逐渐减小,即齿冠逐渐增高的趋势。由于Pararhizomys的牙齿形态多少与竹鼠类的低冠竹鼠Brachyrhizomys和竹鼠属Rhizo-mys及拟速掘鼠属Tachyoryctoides有相似之处,故常被与Brachyrhizomys和Rhizomys一起归入竹鼠科(Rhizom yidae),但该属的牙齿不具中脊和下中脊,褶沟少,一般为2-3个,与上述三属有明显的不同。本文对下门齿所作的切片观察也表明,Pararhizomys的釉质层结构与Brachyrhizomys,Rhizomys和Tachyoryctoides者相差甚远,其内层(PI)明显增厚,釉柱和釉柱间质(IPM)的排列方式存在明显差别。颊齿的冠高、冠面形态,以及下门齿的釉质层结构,似乎都表明了Pararhizomys属不大可能归入竹鼠科或速掘鼠科(Tachyorictoididae)。而下门齿釉质层切片显示了Brachyrhizomys与Rhizomys有着高度相似的釉质结构,进一步证明了两者有较接近的亲缘关系。地理分布上,Pararhizomys与Tachyoryctoides属只发现于古北区,伴生的哺乳动物显示了其可能适应偏冷、干旱的草原环境,而Brachyrhizomys和Rhizomys主要局限于东洋区,共生的哺乳动物多能适应温暖、湿润的树林环境。     

柴达木盆地新近纪犀科化石新材料

近年来在青海柴达木盆地中新世地层内新发现的犀科化石 ,经研究有 3属 3种 ,即Ac erorhinustsaidamensis,Hispanotheriummatritense和Dicerorhinusringstromi,其中后两个种是在这一地区的初次报道。新材料虽较破碎 ,但其发现扩大了H .matritense和D .ringstromi的地理分布范围。更重要的是 ,柴达木动物群原来被确定为晚中新世早期 ,其主要依据是安琪马动物群的残余分子与三趾马动物群共生 ,而此次的发现证明这一地区确实有含H .matritense的中中新世动物群存在。D .ringstromi的发现还证明柴达木盆地有相当于保德动物群时代的晚中新世晚期沉积。     

First identification of Decennatherium Crusafont, 1952 (Mammalia,Ruminantia, Pecora) in the Siwaliks of Pakistan

Previously undescribed remains of a new large giraffid have been identified from the late Miocene Siwaliks Hills of the Potwar Plateau in Pakistan. This taxon is very intimately related to the late Miocene giraffid genus Decennatherium, previously only certainly identified in the Iberian Peninsula, and with some possible remains assigned to Decennatherium crusafonti from Aliäbäd (Iran). The new material collected in the Siwaliks Hills shows a high morphological similarity with the Spanish remains of the genus, especially the early Vallesian Decennatherium pachecoi. Two previously undescribed ossicone fragments from the same area are also described and identified as cf. Decennatherium. These findings represent the easternmost occurrence of the genus Decennatherium and show a late Miocene migration of the genus throughout the Iberian Peninsula and Southern Asia.     

On the origin and development of Artemisia (Asteraceae) in the geological past

Fossil records combined with phytogeographical data reveal that Artemisia L. possibly originated from the existing arid or subarid area of temperate Asia in the mid-Tertiary. The earliest recorded Asteraceae pollen grains are mostly ornamented with medium to long spines, whereas Artemisia L. with short spinules is recorded slightly later. It may be deduced that the latter evolved from the former. Pollen morphological data confirm the coexistence of the two pollen patterns in the subtribe Artemisiinae containing Artemisia L. The development and evolution of the genus suggest three major stages in its history. Artemisia L. did not become abundant until the Middle to Late Miocene in spite of its earlier presence in the Late Palaeogene. Its representation remained low even though there are many records of the genus in the Late Miocene around the world. The Pliocene was an important time for the development of angiosperm herbs including Artemisia L., and the Quaternary has been a major period for the diversification and further worldwide expansion of the genus. The origin and subsequent development of Artemisia L. are considered to be closely related with increased environmental variation, including global climate changes and some important tectonic movements.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 331–336.     

New material of Muntiacinae (Artiodactyla,Mammalia) from the Late Miocene of the northeastern Qinghai-Tibetan Plateau,China

Recent expeditions in the northeastern Qinghai-Tibetan Plateau have produced additional mammalian fossils from the Lower Baodean (equivalent to Vallesian, Late Miocene), and, significantly, a new large and the earliest muntjac, Muntiacus noringenensis sp. nov. It is phylogenetically closer to the Late Pliocene M. fenghoensis and extant M. vuqangensis than the other fossil and extant muntjacs are. The existence of this muntjac and other folivores in the Late Miocene Qaidam Basin suggests a forested period in the basin and the evolution from a forested and humid environment to a desert one today. Such a dramatic evolution in environments is an evidence of the effect of the uplift of the Qinghai-Tibetan Plateau and of its influence on continental climates.     

Italian Cenozoic crocodilians: taxa, timing and palaeobiogeographic implications

Crocodylian remains are collected in 39 fossil-bearing localities but only in seven localities specimens with reliable taxonomic attributions, at least to genus level have been collected. Three species have been reported from the early Lutetian Purga di Bolca site: Pristichampsus cf. Pristichampsus rollinati, Asiatosuchus sp., Hassiacosuchus sp. (=Allognathosuchus sp.). The three crocodilians discovered at Purga di Bolca have been reported also from Geiseltal and Messel (Middle Eocene, Germany). Bolca at that time was part of a Tethysian archipelago and no mammals have been found there till now. Crocodilians and turtles clearly arrived from the European mainland across a marine water barrier. Among the other fossiliferous localities of Veneto, very interesting is the Monte Zuello site, of late Middle Eocene age, yielding a longirostrine crocodilian, Megadontosuchus arduini, a tomistomine species. Tomistomines are known in contemporaneous sediments of both Europe and Africa, but the European forms Dollosuchus and Kentisuchus seem the closest taxa. Remains of Oligocene age have been collected in Veneto and Liguria, but the fossils discovered in the second region are teeth or fragmented bones. The fossil crocodilians of Monteviale (Veneto), of Early Oligocene age, have been assigned to two species but they have been recently all identified as Diplocynodon ratelii, known from several European sites of Late Eocene, Oligocene and Miocene age. This species arrived in the Monteviale area from the European mainland across a narrow sea. Several crocodilian fossils of Miocene age are very fragmentary or represented by isolated teeth. In the Middle and Late Miocene of Sardinia, a well-established species, Tomistoma calaritanum is present. Remains of Tomistoma of the same age have been reported in some localities in Tuscany, Apulia, Sicily and Malta. In the Mediterranean area, the genus is known from European and African sites (of older age). The colonisation of Europe by this genus is the result of a dispersion from Africa (or less probably from Asia). During Late Miocene Sardinia and Tuscany belong to the same palaeobioprovince characterized by the Oreopithecus-Maremmia fauna. In Tuscany, a crocodilian identified as Crocodylus bambolii is present in the late Miocene site of Monte Bamboli. If the generic attribution of this form is correct, its ancestors must have arrived from Africa. Another fossil assemblage of Late Miocene age characterizes the Apulia-Abruzzi palaeobioprovince (Hoplitomeryx-Microtia fauna) and testifies complete isolation between the two palaeobioprovinces. In this last area, remains of Crocodylus sp. have been collected in coastal sandstones at Scontrone (Abruzzi) and in several fissure fillings of Gargano of slightly younger age. The ancestors of this species arrived from Africa while no African elements are present among the mammalian fauna. The dispersion of the genus Crocodylus in the Italian palaeoislands may have taken place once, with allopatric differentiation of the two populations (Tuscany-Sardinia and Apulia-Abruzzi) or twice with independent colonisation of each area.     

Une nouvelle espèce du genre Bohlinia (Mammalia,Giraffidae) du Miocène supérieur de Toros-Menalla,Tchad

New Giraffidae specimens discovered from the Late Miocene fossiliferous sector of Toros-Menalla (northern Chad) are described. These specimens are assigned to Bohlinia, a genus known so far only in some Late Miocene localities of Eurasia. This is the first occurrence of the genus in Africa. Analysis of the material allows us to describe a new species of giraffid, Bohlinia adoumi n. sp., probably close to the Eurasian forms.     

Miocene hyaenids (Hyaenidae; Carnivora) from Siwaliks of Pakistan

Newly discovered Miocene hyaenid specimens, recently collected from the Siwalik Group, are described and discussed. A careful comparison with the known material reveals that these specimens belong to the early hyaenid species Thalassictis cf. T. proava, Ictitherine indet. and Lepthyaena sivalensis. The stratigraphic range of T. proava extended up to the Dhok Pathan Formation (Middle Miocene to Early Pliocene). The stratigraphic range of T. proava comprises the Middle to Late Miocene, with the youngest record in Hasnot, Potwar Plateau in the Siwalik Group. The material is of great interest because Siwalik carnivoran material is rare.     

柴达木盆地晚中新世三趾马化石

Bohlin描述的柴达木动物群中的三趾马材料非常少 ,仅能证明三趾马在这个地点的存在。近年来新的野外考察在这一地区发现了更多的三趾马化石材料 ,至少包括 3个种 ,即Hipparioncf.H .chiai、H .weihoense和H .teilhardi。H .cf.H .chiai和H .weihoense在柴达木盆地的发现进一步证实了晚中新世早期 (保德早期 )动物群在这个地区的存在。柴达木盆地的H .teilhardi基本上与H .cf.H .chiai和H .weihoense产自同一层位 ,其时代也应为保德早期。H .cf.H .chiai和H .weihoense在柴达木盆地的发现为该地区在晚中新世早期为草原型环境的判断提供了更多的证据。H .teilhardi更细长的远端肢骨也是对开阔环境的一种适应性状     

A new duck from the Middle Miocene of Mongolia, with comments on Miocene evolution of ducks

A new relatively large duck, Chenoanas deserta gen. et sp. nov. from the Middle Miocene of the Sharga locality, which is morphologically intermediate between the extant genera Chenonetta and Tachyeres, is described. The diversity of Early and Middle Miocene ducks is discussed. It is noted that some Middle Miocene duck remains are incorrectly referred to the genus Mionetta. The distribution of morphological characters of the humerus in Neogene and extant ducks shows that the present day diversity of ducks apparently results from extinction of some taxa which were formed in the Oligocene-Early Miocene. The distribution of morphological characters in the evolution of diving ducks is evidence that not only the formation of different morphotypes but also so-called ??evolutionary maturation?? of taxa also explains the modern diversity.     

The European Tertiary Neritiliidae (Mollusca, Gastropoda, Neritopsina): indicators of tropical submarine cave environments and freshwater faunas

The oldest freshwater neritiliid, Neritilia bisinuata , is described from the Middle Eocene of the Loire Basin. Another European species, N. neritinoides , ranging from the Lower Oligocene to Lower Miocene (Upper Burdigalian) is recognized; its habitat appears to have been freshwater, but very close to the sea. Two new marine neritiliid species from the Aquitaine Basin are described: Bourdieria favia sp. nov. from the Upper Oligocene and Pisulinella aucoini  sp. nov. from the Lower Miocene. A third undescribed species from the Lower Miocene is referred to the same family and related to Pisulinella . The Oligocene species has a strong spiral sculpture, a character completely absent in previously known neritiliid species. The genus Agapilia , founded on juvenile N. neritinoides and adult Vitta picta , appears to be a junior synonym of the genus Vitta. The associated occurrence of shells of the families Neritiliidae, Neritopsidae and Pickworthiidae (well-known inhabitants of Indo-West Pacific submarine caves) at Peyrère suggest the first occurrence of a characteristic assemblage of dark submarine caves during the Oligocene. Both factorial analysis and relative abundance show that at Peyrère these families are associated with other cryptic fossils (various gastropods, bivalves, Brachiopoda, corals, Annelida). However, there are indications of other submarine cave assemblages in various Cenozoic deposits from the Palaeocene to the middle Miocene.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 447–467.     

New discoveries on the giant hedgehog Deinogalerix from the Miocene of Gargano (Apulia,Italy)

During the Miocene, the Abruzzo/Apulia region (Italy), isolated from the continent, was the theatre of the evolution of a vertebrate ecosystem in insular context. During the Late Miocene, the protagonists of this ecosystem called “Mikrotia fauna” show a high endemic speciation level, presenting spectacular giant and dwarf species of mammals and birds. Deinogalerix is one of the most uncommon forms of this peculiar fauna. It is the largest Galericinae that ever lived. From Gargano (Apulia, Italy) deposits (also called “Terre Rosse”), Deinogalerix has been, since its discovery in the 1970s’, the subject of two publications in which various morphotypes were described. Its presence is also attested at Scontrone (Abruzzo, Italy). In 2005, the new Miocene fissure “Mikrotia 013” (M013) was found in the Gargano area at Cava Dell’Erba by a team of the Università degli studi di Torino. The study of the micromammals, and in particular the presence of an archaic form of Mikrotia, attests that M013 is the oldest fissure known to date. All the Deinogalerix remains come from the smallest morphotype ever found. The morphological features (other than size) allow us to describe the new species Deinogalerix masinii. It presents a large variability of dental and mandibular features expressed in mosaic patterns within the material: the robustness of the upper and lower P3 and P4, and the characteristic trigonid of m1 of the genus; the archaic morphology of the maxilla and the mandible; the abrading pattern of the teeth are unique for Deinogalerix. These new remains allow us to update the knowledge of Deinogalerix. The suit of characters testified that D. masinii nov. sp. is an archaic form on the evolutionary trend toward the most derived representatives of the genus. The affinities of Deinogalerix with the Parasorex group are confirmed, but the time of immigration in Gargano remains debated.     

Miocene pollen record of KC-1 core in the Qaidam Basin,NE Tibetan Plateau and implications for evolution of the East Asian monsoon

Thick Cenozoic deposits in the Qaidam Basin provide great potential for understanding the tectonic history, paleoclimatic changes, and evolution of the East Asian Monsoon. This study examines the pollen record from the KC-1 core for the interval covering the later Early to Late Miocene (18–5 Ma). Thermophilic taxa percentages are high between 18 and 14 Ma and decrease after this time, a pattern which fits well with the Middle Miocene Climatic Optimum (MMCO) between 18 and 14 Ma and global climatic cooling after 14 Ma. During the same period, xerophytic taxa percentages gradually increase and those of the conifers gradually decrease, suggesting an aridification process in the Qaidam region driven by the gradual strengthening of the East Asian winter monsoon (EAWM) and weakening of the East Asian summer monsoon (EASM). The global climate cooling process appears to have driven the climatic development of the Qaidam Basin region throughout the Miocene, but the uplift of the Tibetan Plateau also contributed.     

The spread of grass-dominated habitats in Turkey and surrounding areas during the Cenozoic: Phytolith evidence

The arrival of hipparionine horses in the eastern Mediterranean region around 11 Ma was traditionally thought to mark the simultaneous westward expansion of savanna vegetation across Eurasia. However, recent paleoecological reconstructions based on tooth wear, carbon isotopes, and functional morphology indicate that grasses played a minor role in Late Miocene ecosystems of the eastern Mediterranean, which were more likely dry woodlands or forests. The scarcity of grass macrofossils and pollen in Miocene floras of Europe and Asia Minor has been used to support this interpretation. Based on the combined evidence, it has therefore been suggested that Late Miocene ungulate faunal change in the eastern Mediterranean signals increased aridity and landscape openness, but not necessarily the development of grass-dominated habitats.

To shed new light on the Miocene evolution of eastern Mediterranean ecosystems, we used phytolith assemblages preserved in direct association with faunas as a proxy for paleovegetation structure (grassland vs. forest). We extracted phytoliths and other biogenic silica from sediment samples from well-known Early to Late Miocene ( 20–7 Ma) faunal localities in Greece, Turkey, and Iran. In addition, a Middle Eocene sample from Turkey yielded phytoliths and served as a baseline comparison for vegetation inference.

Phytolith analysis showed that the Middle Eocene assemblage consists of abundant grass phytoliths (grass silica short cells) interpreted as deriving from bambusoid grasses, as well as diverse forest indicator phytoliths from dicotyledonous angiosperms and palms, pointing to the presence of a woodland or forest with abundant bamboos. In contrast, the Miocene assemblages are dominated by diverse silica short cells typical of pooid open-habitat grasses. Forest indicator phytoliths are also present, but are rare in the Late Miocene (9–7 Ma) assemblages. Our analysis of the Miocene grass community composition is consistent with evidence from stable carbon isotopes from paleosols and ungulate tooth enamel, showing that C₄ grasses were rare in the Mediterranean throughout the Miocene. These data indicate that relatively open habitats had become common in Turkey and surrounding areas by at least the Early Miocene ( 20 Ma), > 7 million years before hipparionine horses reached Europe and arid conditions ensued, as judged by faunal data.     

东海陆架西南部台北坳陷新近纪孢粉植物群演替

对东海陆架西南部台北坳陷13口钻井孢粉分析表明,本地区新近纪植物群经历了3个发展阶段,即：1)早中新世早期松科花粉优势期;2)早、中中新世阿丁枫科花粉繁盛期;3)晚中新世至早上新世草本被子植物发展期。各时期孢粉植物群在横向上几乎没有大的区别,表明新近纪整个坳陷内构造与沉积环境及其变迁具有很好的一致性。植物群演替反映古气候从早中新世早期的湿润温凉到早、中中新世的暖湿,最后(晚中新世至早上新世)又趋温凉的变化历程。