New chiropterans from the middle Eocene of Shanghuang (Jiangsu Province,Coastal China): new insight into the dawn horseshoe bats (Rhinolophidae) in Asia

Until recently, the fossil record of Paleogene bats in Asia primarily included extinct families (i.e. ‘Eochiroptera’) from the early Eocene of Vastan in India and from the middle‐late Eocene of the Liguanqiao and Yuanqu basins in central China. Here, we describe a new fauna of Chiroptera from the middle Eocene Shanghuang fissure fillings of China. The fauna includes abundant material referred to a new rhinolophid (Protorhinolophus shanghuangensis gen. and sp. n.), one specimen of a possible rhinopomatid and several indeterminate rhinolophoids. This new bat assemblage constitutes the earliest record of extant families of microbats in Asia. Because it lacks representatives of ‘Eochiroptera’, this Shanghuang bat fauna indicates significant turnover in Asian bat communities. The dental pattern of P. shanghuangensis shows a mosaic of primitive and derived features (‘Eochiroptera’ vs Rhinolophidae dental characteristics), suggesting that this taxon occupies a basal position among the Rhinolophidae. Rhinolophids were already well diversified at the end of the late Eocene in Europe. Interestingly, many dental characteristics of Protorhinolophus are also found in a primitive rhinolophoid taxon, Vaylatsia, from the middle Eocene to late Oligocene of Europe, supporting a close relationship between these taxa. These affinities testify to the widespread Eurasian distribution of rhinolophoids during the Eocene and are consistent with a westward dispersal of the group from eastern Asia to Europe owing to the greater antiquity of Protorhinolophus.     

Species diversity and postcranial anatomy of eocene primates from Shanghuang,China

The middle Eocene Shanghuang fissure‐fillings, located in southern Jiangsu Province in China near the coastal city of Shanghai (Fig. 1 ), contain a remarkably diverse array of fossil primates that provide a unique window into the complex role played by Asia during early primate evolution. 1 Compared to contemporaneous localities in North America or Europe, the ancient primate community sampled at the Shanghuang fissure‐fillings is unique in several ways. Although Shanghuang has some typical Eocene primates (Omomyidae and Adapoidea), it also contains the earliest known members of the Tarsiidae and Anthropoidea (Fig. 2 ), and some new taxa that are not as yet known from elsewhere. It exhibits a large number of primate species, at least 18, most of which are very small (15‐500 g), including some of the smallest primates that have ever been recovered. 2 - 4 © 2012 Wiley Periodicals, Inc.     

Middle Eocene primate tarsals from China: implications for haplorhine evolution

We describe tarsal remains of primates recovered from the Middle Eocene (approximately 45 mya) Shanghuang fissures in southern Jiangsu Province, China. These tarsals document the existence of four higher-level taxa of haplorhine primates and at least two adapid species. The meager and poorly preserved adapid material exhibits some similarities to European adapines like Adapis. The haplorhine primates are divided into two major groups: a "prosimian group" consisting of Tarsiidae and an unnamed group that is anatomically similar to Omomyidae; and an "anthropoid group" consisting of Eosimiidae and an unnamed group of protoanthropoids. The anthropoid tarsals are morphologically transitional between omomyids (or primitive haplorhines) and extant telanthropoids, providing the first postcranial evidence for primates which bridge the prosimian-anthropoid gap. All of the haplorhines are extremely small (most are between 50-100 g), and the deposits contain the smallest euprimates ever documented. The uniqueness of this fauna is further highlighted by the fact that no modern primate community contains as many tiny primates as does the fauna from Shanghuang.     

Squamate reptiles from the middle Eocene of Lissieu (France). A landmark in the middle Eocene of Europe

In Europe, faunas of squamates (lizards and snakes) from the middle Eocene are very poorly known, with the exception of those from the level MP 16 (latest middle Eocene). From the MP 11-MP 15 interval, squamates were previously reported only from Messel (MP 11, earliest middle Eocene) and from the untere and obere Mittelkohle of Geiseltal (MP 12 and MP 13 respectively) in Germany. The present report describes the middle Eocene assemblage of squamates from Lissieu (France), the first fauna reported from the level MP 14. Whereas fossils from Messel and Geiseltal are mostly articulated skeletons, fossils from Lissieu are represented by disarticulated bones; such fossils may be more easily compared to those from other Cenozoic localities, in which bones are almost always disarticulated. The fauna from Lissieu is more diverse than those from the Geiseltal sites and approximately as diverse as that from Messel as they are presently known; it is comprised of 17 distinct taxa. These taxa cannot be all identified to the species or genus level. They belong to iguanids, gekkonids, lacertids, anguids, thecoglossan platynotans, ophidians incertaesedis, boids, ?tropidophiines, “tropidophiids” incertaesedis, booids incertaesedis, and perhaps russellophiids. The fauna includes several new taxa but only a presumed tropidophiine snake may be named on the basis of the available material. The fauna from Lissieu is a mixture of taxa restricted to the middle Eocene and taxa known from older or younger levels. Taxa shared by Lissieu and the few other localities from the middle Eocene of Europe are rare. This fauna from Lissieu represents a stratigraphical landmark for the middle Eocene.     

山东新泰中始新世化石哺乳类新材料

本文记述了采自山东新泰的一批中始新世哺乳动物,并主要地与内蒙大苏尼特右旗呼和勃尔和等地“阿山头组”中的哺乳动物做了比较。新泰动物群似应为中始新世早期。     

内蒙古中始新世一似ameghinornithid鸟类(鸟纲:Cariamae:Ameghinornithidae?)

内蒙古二连盆地额尔登敖包剖面(相当于伊尔丁曼哈组)的中始新世早期地层中新发现的一件标本显示其属于一个与ameghinornithid相似的种,代表了Ameghinomithidae在亚洲的首个记录.新标本具有外踝半圆形,缺失骨质腱桥,伸肌沟外侧具有大而平的结节,以及其他与ameghinomithid和似ameghinomithid鸟类共有的特征.内蒙古标本与欧洲最古老的ameghinomithids记录大体属同一时代(约48 Ma).与同时期内蒙古哺乳动物群主要由亚洲类群组成兼有少量与北美有关的绝灭类群不同,这类鸟类中始新世时在北美缺失,而存在于欧洲和亚洲.加上埃及法尤姆早渐新世地层中发现的似ameghinomithid鸟类,这类已经绝灭的鸟类的地理分布似乎比过去所认为的要广泛得多.     

内蒙古赤峰初头朗早更新世哺乳动物群的新材料及二元相似性分析

出土于内蒙古赤峰初头朗小南山地点的哺乳动物化石新材料有直隶犬(Canis chihliensis),泥河湾披毛犀(Coelodonta nihowanensis),中国长鼻三趾马(Hipparion (Proboscidipparion) sinense),三门马(Equus sanmeniensis),李氏野猪(Sus lydekkeri),湖麂相似种(Muntiacus cf.M.lacustris),山西轴鹿(Axis shansius),布氏真枝角鹿(Eucladoceros boulei),翁氏转角羚羊(Spirocerus cf.S.wongi)及古中华野牛(Bison palaeosinensis).加上此前在初头朗东梁、东村北沟和东村南沟3个地点出土的标本,初头朗一带出土的哺乳动物化石种类增加到30个.初头朗4个地点的哺乳动物化石产自同一层位,属于同一个动物群.根据动物群的二元相似性系数,初头朗动物群在组成上最接近狭义泥河湾动物群.综合Brainerd-Robinson排序结果、动物群古老系数和动物群中绝灭种类的百分比,初头朗动物群的时代介于狭义泥河湾动物群和柳城巨猿洞动物群之间;而根据与相关动物群年龄值的对比,初头朗动物群的年龄值约在1.4-1.6 Ma之间.初头朗动物群中的食肉目种类有1 1个,但大多为小型种类.奇蹄目和偶蹄目种类的总和占动物群总数的一半,大多为体型较大的种类.该动物群中的嫩食者和林柄者指示当时存在一定范围的森林和林地;而动物群中的粗食者及开阔地栖息者指示当时存在面积较大的草原和草甸.初头朗动物群的大部分种类是喜温动物,喜冷种类只有鼠兔和披毛犀,因此当时的气候与现今大致相同.在地理位置上初头朗动物群是我国最靠北和最靠东的早更新世动物群,因此可以推荐为东北地区的早更新世哺乳动物群典型地点.     

First record of non-marine ostracods from the Paleogene “hamadian deposits” of Méridja area,west of Bechar (southwestern Algeria)

A new non-marine ostracod fauna from the Paleogene “hamadian deposits” outcropping west of Bechar (southwestern Algeria) has been recovered from lacustrine to fluvial deposits of the Oued Méridja section and fluvial deposits on the southern edge of the Hamada de Méridja section. Recently, these sections have been dated as late Thanetian – early Ypresian (latest Paleocene to earliest Eocene) and Ypresian – earliest Lutetian (early to earliest middle Eocene), respectively, based on charophytes. The associated ostracod fauna recovered consists of relatively mostly moderately to badly preserved specimens and comprises 14 taxa, none of which could be identified to species level in view of its poor state of preservation; we have nevertheless been able to identify and describe the following taxa: Herpetocypris sp., Cyprinotus? sp., Heterocypris? sp. 1 and sp. 2, Cypris? sp., Ilyocypris sp., Cytheroidea indet. sp. 1 and sp. 2, Limnocytheridae indet. sp. 1, Cypridoidea indet. sp. 1, Cyprididae indet. sp. 1, and Ostracoda indet. sp. 1, 2 and 3. Only Heterocypris sp. 1 occurs in both sections. Although the fauna can as yet not be related to the few other contemporaneous faunas reported from the wider palaeogeographic area, it adds important new information to our poor knowledge on Eocene non-marine ostracods in North Africa and southern Europe. The Méridja sections and area are promising regarding the discovery of more, better preserved material and further studies, and one main limitation to the correlation of the fauna is the hitherto insufficient taxonomic knowledge on many faunal elements of Eocene non-marine ostracods to which our section contributes considerably.     

An assemblage of early Oligocene lizards (Squamata) from the locality of Boutersem (Belgium), with comments on the Eocene–Oligocene transition

A diverse, new lizard assemblage from the early Oligocene of Belgium is described. The Boutersem railway local fauna is the most species‐rich lizard assemblage yet reported from the European early Oligocene. Four lizard taxa are present: Lacertidae, Anguidae, Scincoidea and Platynota. One new species is described, Folisaurus boutersemensis sp. nov . This fauna provides new insight into the profound turnover that took place during the Eocene/Oligocene boundary in Europe. The new fauna confirms a marked decrease in diversity across the Eocene/Oligocene boundary. Two groups encountered in the European late Eocene became extinct (Iguanidae*, Glyptosaurinae). Estimates of species‐level extinctions range up to 80%. These estimates include members of virtually all the families present in the late Eocene. The relative importance of climate change and biotic interactions in controlling this pattern is discussed, and negative interactions between lizards and new carnivorous mammals are favoured. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 148–170.     

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.     

Significance of primate petrosal from Middle Eocene fissure-fillings at Shanghuang, Jiangsu Province, People's Republic of China

An isolated petrosal bone belonging to a diminutive primate is reported from Middle Eocene fissure-fills near Shanghuang (southern Jiangsu Province, People's Republic of China), the type locality of several newly described primates (Eosimias sinensis, a basal anthropoid; Adapoides troglodytes, a basal adapinan; Tarsius eocaenus, a congener of extant tarsiers; and Macrotarsius macrorhysis, the first Asian representative of an otherwise exclusively North American genus). Because of its fragmentary condition and unique combination of characters, the Shanghuang petrosal cannot be assigned unambiguously to any of the Shanghuang primate taxa known from dental remains. However, the possibility that the petrosal represents either an adapid or a tarsiid can be dismissed because it lacks defining basicranial apomorphines of these groups. By contrast, the element does present arterial features consistent with its being haplorhine. Deciding between the likeliest candidates for its allocation—Omomyidae and Eosimiidae—is difficult, in part because it is not known what (or even whether) basicranial characters can be used to distinguish these clades. If the Shanghuang petrosal is that of an cosimiid, as both direct and indirect evidence appears to indicate, the following implications emerge: (1) as long suspected on other grounds, anthropoids share a closer evolutionary history with Omomyidae (and Tarsiiformes) than they do with Adapidae (and Strepsirhini); (2) the specialised basicranial anatomy of extant anthropoids and their immediate cladistic relatives is derived from a primitive precursor whose otic morphology was like that of omomyids in most known respects; (3) the evolution of the defining dental and basicranial apomorphies of extant Anthropoidea has been distinctly mosaic in pattern.     

山西垣曲河堤组寨里段的低等四足类动物群

本文记述了我国晚始新世的一低等四足类动物群,包括三种蜥蜴和两种鳄类.其中的宽额半鳄蜥(Hemishinisaurus latifrons gen. et sp. nov.)是我国首次发现的异蜥科化石;垣曲响蜥(Tinosaurus yuanquensis sp. nov.)是响蜥属在我国的最晚代表.     

河南许昌灵井旧石器遗址哺乳动物群的性质及时代探讨

系统记述了在近两年从河南省许昌市灵井旧石器遗址出土的食肉目、长鼻目和奇蹄目化石:宽吻灵猫相似种(Viverra cf. zibetha)、熊未定种(Ursus sp.)、中国硕鬣狗相似种(Pachycrocuta cf. sinensis)、古菱齿象未定种(Palaeoloxodon sp.)、披毛犀(Coelodonta antiquitatis)、梅氏犀(Dicerorhinus mercki)、普通马(Equusca ballus)和蒙古野驴(Equus hemionus),并对灵井哺乳动物群的性质和年代进行了探讨,指出灵井动物群含有的北方种类较多,与北方区的很多动物群可以比较,属于北方区动物群,其生态环境为以平原草地为主,镶嵌有零散的森林,属北温带半湿润-半干旱大陆性季风气候,年平均温度似低于现在。灵井动物群中绝灭种类占44.4%,其时代应为晚更新世早期,其绝对年龄与许家窑接近,约在100kaBP。     

Fossil Evidence and the Origin of Bats

The phylogenetic and geographic origins of bats (Chiroptera) remain unknown. The earliest confirmed records of bats date from the early Eocene (approximately 51 Ma) in North America with other early Eocene bat taxa also being represented from Europe, Africa, and Australia. Where known, skeletons of these early taxa indicate that many of the anatomical specializations characteristic of bats had already been achieved by the early Eocene, including forelimb and manus elongation in conjunction with structural changes in the pectoral skeleton, hind limb reorientation, and the presence of rudimentary echolocating abilities. By the middle Eocene, the diversification of bats was well underway with many modern families being represented among fossil forms. A new phylogenetic analysis indicates that several early fossil bats are consecutive sister taxa to the extant crown group (including megabats), and suggests a single origin for the order, at least by the late Paleocene. Although morphological studies have long placed bats in the Grandorder Archonta, (along with primates dermopterans, and tree shrews), recent molecular studies have refuted this hypothesis, instead strongly supporting placement of bats in Laurasiatheria. Primitively, proto-bats were likely insectivorous, under-branch hangers and elementary gliders that exploited terminal branch habitats. Recent work has indicated that a number of other mammalian groups began to exploit similar arboreal, terminal branch habitats in the Paleocene, including multituberculates, eulipotyphlans, dermopterans, and plesiadapiforms. This may offer an ecological explanation for morphological convergences that led to the erroneous inclusion of bats within Archonta: ancestral archontan groups as well as proto-bats apparently were exploiting similar arboreal habitats, which may have led to concurrent development of homoplasic morphological attributes.     

Western Amazonia as a Hotspot of Mammalian Biodiversity Throughout the Cenozoic

A state-of-the-art review of the Cenozoic fossil record from Western Amazonia is provided, based on literature and new data (regarding Paleogene native ungulates). It allows summarizing the evolution and dynamics of middle Eocene–Holocene mammalian guilds, at the level of species, families, and orders. Major gaps in the Western Amazonian mammal record occur in the pre-Lutetian and early Miocene intervals, and in the Pliocene epoch. Twenty-three orders, 89 families, and 320 species are recognized in the fossil record, widely dominated by eutherians from the middle Eocene onward. Probable Allotheria (Gondwanatheria) occur only in the earliest interval, whereas Metatheria and Eutheria are conspicuous components of any assemblage. Taxonomic diversity was probably fairly constant at the ordinal level (~12–14 orders in each time slice considered) and much more variable in terms of family and species richness: if most intervals are characterized by 40–50 co-occurring species and 19–31 co-occurring families, the early Miocene period illustrates a depauperate fauna (21 species, 17 families), strongly contrasting with the late Miocene climactic guild (82 species, 38 families). Recent mammalian taxonomic diversity from Western Amazonia (12 orders, 37 families, and 286 species) is at odds with all past intervals, as it encompasses only three orders of South American origin (Didelphimorphia, Cingulata, and Pilosa) but four North American immigrant orders (Artiodactyla, Perissodactyla, Carnivora, and Lagomorpha). In terms of taxonomic diversity, recent mammalian guilds are fully dominated by small-sized taxa (Chiroptera, Rodentia, and Primates). This overview also confirms the scarcity of large mammalian flesh-eaters in ancient Neotropical mammalian assemblages. The pattern and the timing of mammalian dispersals from northern landmasses into Western Amazonia are not elucidated yet.     

HIRNANTIAN (LATEST ORDOVICIAN) GRAPTOLITES FROM THE UPPER YANGTZE REGION, CHINA

Abstract:  The Upper Yangtze region yields a Hirnantian (latest Ordovician) graptolite fauna that includes 41 species assigned to 13 genera. This fauna is particularly important for understanding the Late Ordovician mass extinction event because it is the most diverse known from this interval. In addition, it records the survival, well into the Hirnantian, of many taxa of the Dicranograptidae-Diplograptidae-Orthograptidae (DDO) fauna, which was previously regarded as having gone extinct at the beginning of the Hirnantian. Taxa exhibiting six different astogenetic patterns, including taxa with reclined stipes, scandent, biserial, full-periderm and 'archiretiolitid' rhabdosome forms occur in the lower Normalograptus extraordinarius-N. ojsuensis Biozone. In contrast, in the upper N. persculptus Biozone only four genera remain, all but one of which are Normalograptidae: scandent and biserial taxa with Pattern H astogeny. Normalograptids are the dominant form of the succeeding, lower Rhuddanian, faunas. The Yangtze faunas also document the early expansion of normalograptids coeval with the decline of the DDO fauna. Many previously identified species considered endemic to China have been synonymized; 24 of the 41 species recorded here have been recognized elsewhere. No new taxa are described.     

辽宁朝阳龙城马山洞哺乳动物群的性质及时代探讨

系统记述了在2007年从辽宁省朝阳市龙城区马山洞出土的奇蹄目和偶蹄目化石:普氏野马(Equus przewalskii)、马鹿(Cervus(E.)elaphus)、普氏原羚(Proca praprzewalskii)、恰克图转角羚羊(Spirocerus kiakhtensis)、裴氏转角羚羊(Spirocerus peii)和岩羊未定种(Pseudois sp.)。并对马山洞哺乳动物群的性质和年代进行了探讨, 指出马山洞动物群含有的北方种类较多, 与北方区的很多动物群可以比较, 属于典型的北方区动物群。其生态环境为以山地原草地为主, 镶嵌有零散的森林, 属北温带半湿润-半干旱大陆性季风气候, 年平均温度似低于现在。马山洞动物群中绝灭种类不多, 主要为晚更新世的类型, 在组成上与古龙山、小孤山和阎家岗这3地的动物群最相似, 其时代似应为晚更新世中晚期, 其绝对年龄估计在5ka左右。     

New primate hind limb elements from the middle Eocene of China

The continued washing, sorting, and identification of middle Eocene (45 Mya) primates from the Shanghuang fissure-fillings (Jiangsu Province, China) have produced additional hind limb elements. All are isolated elements. The strepsirhine hind limb elements include a first metatarsal and a talus, which are appropriate in size and morphology to pertain to Adapoides troglodytes. Adapoides is interpreted as a quadrupedal-climbing (nonleaping) primate with similarities to living lorises and the fossil primate Adapis. The haplorhine hind limb elements are estimated to span a range of adult body sizes from tiny (17 g) to small (200 g). Included among the new sample of haplorhine hind limb specimens is the smallest primate talus reported thus far. These new postcranial specimens expand our understanding of early haplorhine hind limb anatomy and demonstrate additional similarities between Shanghuang eosimiids and other anthropoids.     

甘肃灵台上新世哺乳动物化石埋藏学

灵台任家坡动物群年龄组成以老年、幼年个体为主,显示动物正常死亡条件下的骨骼富集积累过程。由于动物死亡与掩埋之间时限的长短不同导致个体间风化痕迹、关节分解、破损程度不同。化石具有明显的定向性及分选性,经过季节性水流短距离搬运和较弱的分选作用。化石的埋藏特点与泛滥平原骨骼富集有较多的相似性,但沉积物组成差别很大,暗示中国红粘土物源的特殊性。