Middle Miocene Cephalotaxus (Taxaceae) from Yunnan,Southwest China,and its implications to taxonomy and evolution of the genus

The infrageneric taxonomy of the genus Cephalotaxus Siebold et Zuccarini ex Endlicher (Taxaceae) is controversial as morphological characters have been doubted to change coherently among species. Although the epidermal features are considered as important characters in the taxonomy of conifers, they have not been systematically studied in this genus. Neogene foliage shoots with well-preserved epidermal structure not only can provide systematic links to the extant species, but also have implications for the taxonomical definition of the extant species by their epidermal features. In this paper, the morphologies of leafy shoots combining epidermal structure of a Neogene Cephalotaxus, which was collected from the middle Miocene of Southwest China, and ten extant species (variants) were studied. Morphology as well as epidermal structures was compared among species (variants) in Cephalotaxus. Based on the comparisons, the taxonomical definition of extant species in genus Cephalotaxus are discussed. Our results indicate that leaf morphological characters in combination with the features of epidermal structure can distinguish species in genus Cephalotaxus. The fossil foliage shoots from the middle Miocene of Southwest China were assigned to a new species, C. maguanensis n. sp., which shows close affinities to the extant C. hainanensis. On the basis of leaf and epidermal information, we suggest there are seven extant species in Cephalotaxus. With global cooling and the uplift of the Qinghai-Tibet Plateau since the middle Miocene, the cooling climate induced the differentiation and southward expansions of C. maguanensis, and its extant offspring, C. hainanensis, is distributed to Hainan Island and the Indo-Chinese peninsula.     

Fossil leaves of Populus from the Middle Miocene of Yunnan,SW China

Today southern Yunnan, SW China, has a tropical or subtropical climate and seasonal rainforests. In the past, some temperate elements were also present. In this paper, a new species of Populus is reported from the Middle Miocene deposits in Zhenyuan. Its leaves are ovate or ovate-suborbicular, with serrate margins. They have a shallowly cordate to cuneate base without glands, short acuminate apex, and salicoid teeth with spherical glands. The veins are glabrous but unicellular hair bases occur on the lower epidermis of the lamina. Stomata are confined to the lower epidermis. The presence of Populus in the Middle Miocene of the region indicates an expansion of the genus into low-latitude Asia in the late Cenozoic and a more complicated history of vegetational change in southern Yunnan than has so far been assumed.     

New material of Hispanotherium matritense (Rhinocerotidae, Perissodactyla) from Laogou of Hezheng County (Gansu, China), with special reference to the Chinese Middle Miocene elasmotheres

New rhinocerotid teeth from Laogou in Hezheng (Gansu, China) are described and identified as Hispanotherium matritense. Comparing the other Chinese Middle Miocene elasmotheres with material of H. matritense from Iberian localities and Laogou, we confirm the synonymy proposed previously for the species Hispanotherium lintungensis, Tesselodon fangxianensis, Caementodon tongxinensis and Huaqingtherium qiui. The larger size of Hispanotherium tungurense proves that it is a different species from H. matritense. The age of Laogou fauna corresponds to MN6. Paleoenvironments of H. matritense in China include rivers and lakes, which are somewhat different from the arid environment in the Iberian Peninsula. The origin of the Hispanotherium lineage may be in southwestern Europe. After that, it dispersed eastward to Asia.     

云南中新世杉科化石木研究

利用光学显微镜和扫描电镜首次对云南省楚雄州晚中新世石灰坝组石鼓村层的钙化木材进行了解决学研究。鉴定出两种类型的木材。柳杉型落羽杉型木（Taxodioxylon cryptomeripsoides Schonfeld)和杉木型落羽杉型木（Taxo-dioxylon cunninghamioides Watari)。二者分别与现代柳杉属和杉木属具有最接近的亲缘关系。根据这两种杉科化石木现存最近亲缘种的生态环境,并综合其他资料,推测该地区在晚中新世为温暖湿润的亚热带气候环境。     

云南中新世杉科化石木研究

利用光学显微镜和扫描电镜首次对云南省楚雄州晚中新世石灰坝组石鼓村层的钙化木材进行了解剖学研究.鉴定出两种类型的木材:柳杉型落羽杉型木(Taxodioxylon cryptomeripsoides Schonfeld)和杉木型落羽杉型木(Taxodioxylon cunninghamioides Watari).二者分别与现代柳杉属和杉木属具有最接近的亲缘关系.根据这两种杉科化石木现存最近亲缘种的生态环境,并综合其他资料,推测该地区在晚中新世为温暖湿润的亚热带气候环境.     

Firmiana (Malvaceae: Sterculioideae) fruits from the Upper Miocene of Yunnan,Southwest China

A new species of Firmiana Marsili is recognized and described based on capsules from the Late Miocene Lincang flora of Yunnan, China. Firmiana yunnanensis nov. sp. is represented by opened, dehiscent, membranous follicles with a weak, sinuous midvein, obtuse secondary veins and thick marginal vein. The follicles conform in shape and venation with those of extant species of Firmiana, a genus now distributed in eastern Africa, Indo-Malaysia, and eastern and southeastern Asia. This fossil species occurred within the native distribution of extant Firmiana species. It provides new information on fossil history, paleoecology, as well as on the plant diversity of the Lincang flora.     

Quantitative reconstruction of palaeoclimate from the Middle Miocene Shanwang flora, eastern China

The fossil leaf flora from the Shanwang Formation, Linqu County, Shandong Province, eastern China, is diverse, abundant, well preserved and discretely spaced, all of which make it suitable for the recognition and application of leaf physiognomic characters. This article is the first study of Chinese Tertiary floras using the CLAMP (Climate Leaf Analysis Multivariate Program) approach. Numerous bedding planes yield leaf assemblages through over 20 m thickness of diatomaceous shales within the Middle Miocene Shanwang Formation. The diatomaceous shales, containing abundant fossils, have been divided into 19 units. A working platform (3×3 m) in the Diatomite Quarry of Shanwang was studied for about 3 months in 1998 and as many fossils as possible were recorded layer by layer. This collecting method, used for the first time in the history of Shanwang research, enabled us to record the presence or absence of megafossils in distinct units so that we could investigate leaf physiognomic characteristics and palaeoclimate for each time interval. Leaf fossils from six units (5, 6, 7, 13, 15 and 16) have been used to estimate palaeoclimate parameters of the Shanwang area using the CLAMP 3B dataset. The predicted climate is that of a lakeside setting, at over 1000 m above sea level and within the 17–15-Ma time interval. Meteorological factors have been estimated using the CLAMP approach, including mean annual temperature (MAT; 9.5–11.2°C), warmest month mean temperature (20.2–22.5°C), growing season length (5.9–6.8 months), mean growing season precipitation (83–165 cm) and relative humidity (73–79%). MAT estimates from leaf margin analysis (LMA) range from 12.5 to 15.2°C, which are a maximum of 4° higher than those from our CLAMP analyses and from previous results of LMA studies of the Shanwang flora (10–13°C). Nearest living relatives (NLR) of other biota (pollen, fish, insects, diatoms, microthyriaceous fungi, etc.) indicate warm and moist temperate to subtropical conditions in the Shanwang area during the Miocene. CLAMP predictions are similar to the climate predicted from NLRs although the MAT prediction is a few degrees cooler. The fact that the Shanwang leaf flora is from a lacustrine sequence may have biased the CLAMP analysis towards cooler (and possibly wetter) climates. The modern Yangtze River valley is a reasonable environmental analogue for the Shanwang Middle Miocene, though Shanwang may have had cooler summers, cooler MAT and less seasonality in rainfall. The climate was essentially constant during deposition of the sequence studied, with possible minor variations in precipitation.     

Two new fossil woods from the early Miocene of Kutch,Gujarat, India and their significance

The Kutch region of western India (Gujarat State) is today arid to semiarid and characterised by mostly ephemeral streams which carry water during the monsoon. The uneven distribution of rainfall and disturbed topography are the result of climate change during the Cenozoic period. Two fossil woods, namely Bauhinium palaeomalabaricum Prakash and Prasad (Fabaceae) and Ebenoxylon indicum Ghosh and Kazmi (Ebenaceae), are described from Kutch in order to provide insights into the palaeovegetation and palaeoclimate. Because the modern representatives of the present and previously described taxa from the same horizon are thermophilic in nature and grow in evergreen to deciduous forests, a warm and humid climate is interpreted. Furthermore, the finding of some mangrove taxa in the assemblage indicates the lagoonal to intertidal environment at the time of deposition.     

A new species of the rhinoceros Alicornops from the Middle Miocene of the Linxia Basin, Gansu, China

A new species of the genus Alicornops, A. laogouense , from Laogou in Hezheng County, Linxia Basin, Gansu, China, is described. It is mid-sized in the subfamily Aceratheriinae, but is the largest known species of Alicornops . It represents the first discovery of the genus in Asia. The Middle Miocene fauna bearing A. laogouense is contemporaneous with the Dingjiaergou fauna of Tongxin, Ningxia, China, and its age corresponds to MN 6 in Europe. The discovery of A. laogouense in the Linxia Basin implies that Alicornops dispersed from Europe to Asia. During MN 6 times it was relatively widespread throughout Eurasia. Alicornops laogouense lived in open woodland rich in lakes and rivers.     

红杉属植物在中国云南中新世首次发现

化石标本采自中国云南省中部楚雄市吕合煤矿的石鼓村层 ,该层为灰色至浅灰色细砂岩至粗砂岩、粉砂质粘土层和褐煤层 ,地质时代属中新世 ① 。石鼓村层的孢粉组合以被子植物占绝对优势(90 % ) ,其中热带、亚热带植物成分类型较多 ,如金缕梅科、棕榈科等 ,温带植物成分类型较少 ,如榆科、桦木科等 ;裸子植物花粉占整个孢粉组合的 9% ,主要是松科和杉科花粉 ;蕨类植物孢子仅占整个孢粉组合的 1% ,以水龙骨单缝孢为主。本次发现的标本小枝长约 2 .8ｃｍ ,宽约 1.6ｃｍ ,小枝基部稍膨大。叶螺旋状排列 ,基部扭转 ,在同一平面排成 2列 ,以 30～ 70…     

Middle Jurassic Ginkgo leaves from the Daohugou area,Inner Mongolia,China and their implication for palaeo-CO2 reconstruction

Well-preserved Ginkgo leaves with cuticle were collected from the Middle Jurassic Jiulongshan Formation in the Daohugou area, Inner Mongolia, China, which form the common elements in the Daohugou flora. Three new species of Ginkgo are recognized and their leaf morphology and cuticular structure are described for the first time. Two different pCO₂ proxy models were applied to reconstruct palaeo-atmospheric CO₂ concentration by using these new Ginkgo material. NLE (nearest living equivalent) suggests a semi-quantitative pCO₂ estimate as 839 ± 99 ppmv with Carboniferous standardization and 419 ± 49 ppmv with Recent standardization. While Barclay’s revised SI-pCO₂ regression suggests the estimates as 405 ± 71 ppmv. The results show that the pCO₂ estimate is higher than today’s atmospheric CO₂ concentration but lower than most of the other results on the Jurassic Period. Combined with the floristic composition, the climate at 165 Ma in the Daohugou area is inferred to be warm temperate.     

Anatomical and developmental study of petrified Quercus (Fagaceae) fruits from the Middle Miocene, Yakima Canyon, Washington, USA

The first reported petrified acorns to show internal anatomical structure are here described from Middle Miocene (～15.6 million years old) chert of the Columbia River Basalt Group in Yakima Canyon, Washington. Quercus hiholensis Borgardt et Pigg sp. nov. is described from anatomical and morphological fruit features, as well as a little recognized anatomical feature, the umbilical complex. Acorns, each comprising a nut and its cupule, are up to 15.3 mm long and 18.8 mm wide with helically arranged, imbricate, tuberculate cupule scales. They show basal aborted ovules, short styles, broad stigmas, and lack grooves in their cotyledons. These characters and the developmental pattern seen in these fossil acorns demonstrate that Q. hiholensis conforms to genus Quercus (Fagaceae), subgenus Quercus, section Quercus (the white oaks). The correspondence of Q. hiholensis to the modern section Quercus reveals that the derived floral and fruit characters that distinguish section Quercus within the genus had evolved by the Middle Miocene.     

First Thyreophoran Type Tracks from the Middle Jurassic Chuanjie Formation of Yunnan Province,China

Two tridactyl footprints from the Chuanjie Formation (Middle Jurassic) of Yunnan Province, China are morphological characteristics of thyreophoran tracks. They show some similarities to Shenmuichnus, known from the Early Jurassic strata of both Shaanxi and Yunnan provinces, but are somewhat larger, thereby resembling the ichnogenus Stegopodus. Based on their general morphology and size being congruent with this ichnogenus, they are tentatively assigned here to cf. Stegopodus. This is the fourth report of large ornithischian (probably thyreophoran) tracks from the Lower-Middle Jurassic of China that indicates relatively large trackmakers that were likely to be taxonomically distinct from much smaller and gracile Anomoepus trackmakers, also of ornithischian affinity. The larger tracks indicate a hitherto unreported abundance, size range and diversity of track types attributed to this group. The parallel orientation of the two best preserved trackways may indicate gregariousness.     

我国西南植物区系的分异: 椿榆属和臭椿属化石的启示

滇西北‒西藏和滇东南‒桂西是我国西南植物区系中两个面貌和谱系年龄差异最为鲜明的区域, 分别以区系年轻、草本植物和区系古老、木本植物为特征。其中一个饶为有趣的问题是这两个地区的区系分异现象是如何演变形成的? 古植物学研究能为回答这一问题提供重要证据。本文报道分别发现于滇西北‒西藏和滇东南‒桂西的椿榆属(Cedrelospermum)和臭椿属(Ailanthus)化石新纪录。结合已有研究, 这两个类群在我国西南的化石历史为: 椿榆属见于西藏中始新世至晚渐新世(西藏椿榆C. tibetica)及滇东南晚始新世(亚洲椿榆C. asiaticum)地层, 而臭椿属化石见于西藏中始新世至晚始新世(大果臭椿A. maximus)及滇东南晚始新世至早渐新世(古臭椿A. confucii)地层。以上古近纪滇西北‒西藏和滇东南‒桂西植物区系在属级水平上的相似性表明这两个地区之间曾经存在区系交流、类群组成相近。然而, 在种级水平上, 自晚始新世以来其椿榆属和臭椿属植物又各不相同, 暗示着以上两个区系可能在晚始新世或更早已经由于环境分异等因素而向着不同的方向发展。现在椿榆属和臭椿属都已经从西藏消失了, 取而代之的是总体上更为年轻的草本植物区系。与此不同, 臭椿属在滇东南‒桂西延续到了现在, 体现了该地区区系的古老性。本研究为滇西北‒西藏和滇东南‒桂西植物区系在地质历史时期的相似性及其分异的形成提供了化石证据。     

Fossil record of a silurid catfish from the Middle Miocene Sanuki Group of Ohkawa, Kagawa Prefecture, Japan

A fossil specimen identified as a silurid catfish (92.5 mm SL) was collected from the Middle Miocene Sanuki Group (15.8±0.9 Ma) of Ohkawa, Kagawa Prefecture, Japan. This is the oldest certain record of a silurid catfish. Morphological characteristics of the specimen are as follows: 52 or 53 vertebrae, 61 or 62 anal fin rays, a stout, short spine (11%SL) and 10 or more soft rays of the pectoral fin, 8 or more pelvic fin rays, 8 or more branchiostegal rays, head length of ca. 24%SL, and most probably not forked caudal fin. The fossil specimen is clearly distinct from three extant silurids in Japan, and probably belongs to an undescribed form considering its geological age.     

Late Miocene Microstonyx remains (Suidae, Mammalia) from Northern China

Several large suid cranial remains attributed to Microstonyx major are part of a new Hipparion Fauna collection from the Hezheng area, Northern China. The new material confirms the presence of Microstonyx in the late Miocene of the area. The Chinese form belongs to a small-sized eastern population with reduced premolar row and clear sexual bimodality. Statistical comparison shows that Microstonyx major was a polymorphic species and reinforces recognition of Hippopotamodon as a separate genus, defined by relatively stout premolars resulting from a different underlying pattern of allometric growth. The presence of Microstonyx in North China and the distinct suid assemblage that lived there suggest biogeographic connections between Northern China and Western Eurasia in contrast to isolation from Southern China and the Indian subcontinent. The suid fauna of the late Miocene of Northern China seems to have been restricted to the later, more humid phase represented by the Red Clay faunas.     

First record of Eretmophyllum (Ginkgoales) with well-preserved cuticle from the Middle Jurassic of the Ordos Basin,Inner Mongolia,China

The genus Eretmophyllum, a characteristic and widespread taxon of the order Ginkgoales, is restricted in the Mesozoic floras of Eurasia. In China, although some specimens were assigned to Eretmophyllum based on gross leaf morphology, none have epidermal structure, which is essential for species delimitation within this genus. In this paper, we describe a new species, Eretmophyllum neimengguensis n. sp., from the Middle Jurassic Yan’an Formation of the Ordos Basin, China. This is the first record of the genus Eretmophyllum (Ginkgoales) from the Ordos Basin as well as from China based upon the gross leaf morphology and the epidermal structure. The new record significantly extends the stratigraphic and geographic distribution of Eretmophyllum in China and Eurasia. It also improves our understanding of its leaf morphology, epidermal structure, and past diversity. The taxonomic position and the stratigraphic and geographical range of the genus Eretmophyllum are discussed. Briefly, leaf morphological and epidermal characters of the new species, associated plant assemblages, and sedimentology indicate that the regional climate of the Ordos Basin was warm and humid with seasonal temperature and precipitation fluctuations in warm-temperate zone during the Middle Jurassic. Furthermore, given the relationship between the distribution/diversity of the genus and the climate, the genus Eretmophyllum may be confined to the warm-temperature climate, is intolerant of heat and frost, and can be considered as an indicator of humid and warm climatic conditions.     

Merychippus (Mammalia, Perissodactyla, Equidae) from the Middle Miocene of state of Oaxaca, southeastern Mexico

Mexican material referable to Merychippus from two localities in eastern Oaxaca was described first nearly 50 years ago. Subsequent work there and in Central Oaxaca, spanning some 30 years, has allowed to establish the detail stratigraphy in both regions, and assembled a collection of merychippine material from the Matatlán (Central Oaxaca) and El Camarón (eastern Oaxaca) Formations, both K-Ar dated ~15 Ma (late early Barstovian). Detailed taxonomic analysis of this collection indicate the presence of two subhypsodont horse species referable to “Merychippus” cf. “M.” primus and “M.” cf. “M.” sejunctus in both regions. These records document the coexistence in tropical southern North America of basal and hipparionine affinity merychippine grade species, and provide a glimpse in to the diversity of subhypsodont equids in this region.     

The first fossil Microsoroid fern (Palaeosorum ellipticum gen. et sp. nov.) from the middle Miocene of Yunnan,SW China

The microsoroid ferns make up a large group in the family Polypodiaceae, easily characterized by the combination of at least partly clathrate rhizome scales and anastomosing reticulate venation. The diversification of this clade is poorly known, both because of unresolved generic delimitations and the lack of fossil records. In this work, we describe the first microsoroid macrofossil: Palaeosorum ellipticum sp. nov. F. M. B. Jacques & Z. K. Zhou. The fossil specimen, represented by part and counterpart, was found in the middle Miocene sediment of the Dajie Formation in Ailsohan, central Yunnan, southwest China. Epiphytic ferns belonging to Drynariaare found in different locations in Yunnan during the Pliocene. The fossil described in this study deepens the occurrence of epiphytic ferns in Yunnan back to the middle Miocene. It demonstrates that forests with complex ecological relationships have existed in southwest China for more than 10 million years.     

New fossil leaves and fruits of Lauraceae from the Middle Miocene of Fujian,southeastern China differentiated using a cluster analysis

The fossil record of Lauraceae can be traced back to the Early Cretaceous of eastern Asia based on fossil flowers. Here, we refer a number of new occurrences of leaf and fruit fossils of Lauraceae from the Middle Miocene of Zhangpu, Fujian, China, to seven species. These data provide evidence supporting the fact that a diverse subtropical, or tropical, Lauraceae-dominated evergreen forest surrounded this region 15 million years ago (Mya). The Lauraceae fossils presented in this paper provide evidence for the evolution of this group as well as new materials that enable the study of the Fujian Province Neogene flora. The fossils described in this paper fill in the gaps in studies about Lauraceae pollen in the Middle Miocene from Fotan, Fujian, China. In addition, these fossils also enrich the Middle Miocene fossil records of Lauraceae in eastern Asia, especially improving the study of the macrostructures and reproductive organs of fossil Lauraceae from southern China. The similarity between fossil and modern fruits shows that during the Middle Miocene the fruit morphological of Lauraceae have changed very little. We also identify families where the fossils we report belong to their closest relatives and can be used to reconstruct the paleoenvironment of Fujian in the Middle Miocene.