From shallow to deep water: an ecological study of the Hirnantia brachiopod Fauna (Late Ordovician) and its global implications

A large brachiopod fauna collected through the upper Wanyaoshu Formation (middle Hirnantian), West Yunnan, China displays, for the first time, a vertical transition within the Hirnantia fauna. The inner to mid-shelf Fardenia–Hirnantia association segues into the deeper water Aegiromena–Anisopleurella association during the Metabolograptus persculptus marine transgression. This transition parallels the pattern apparent in onshore–offshore gradients within the Hirnantian, imitating Walther’s Law for facies, instead predicting the seaward transitions between community types within a single transgressive sequence. In addition, large-sized brachiopods such as Hirnantia and Eostropheodonta become rarer and small-sized brachiopods, for example, Aegiromena and Anisopleurella more common. It is a rare opportunity to view transformations in community composition and structure within the Hirnantia fauna together with changes in both the size of individuals and populations (from large to small) during the transgression.     

The latest Ordovician Hirnantia brachiopod fauna of Myanmar: Significance of new data from the Mandalay Region

A new, latest Ordovician brachiopod fauna is systematically described from the Hwe Mawng Purple Shale Member (Hirnantian) of the Naungkangyi Group of the Pa-thin area, Mandalay Region, Myanmar, revealing one of the most diverse representatives of the typical Hirnantia Fauna. Rhynchonelliformean, craniiformean and linguliformean brachiopods studied belong to 23 genera and a few indeterminate taxa, among which the most abundant genus Kinnella, along with the four common genera (Paromalomena, Pseudopholidops, Fardenia, and Dalmanella), and other genera (such as Cliftonia, Draborthis, Hindella, Hirnantia, Leptaena, Mirorthis, Plectothyrella, Skenidioides, and Xenocrania) are recorded for the first time. This high diversity Hirnantia Fauna represents an ecological differentiation within the benthos of the Sibumasu Terrane during the end Ordovician global crisis. The palaeoeco-unit named herein as the Kinnella-Paromalomena Association is assigned to lower BA3. This paper includes a taxonomic revision with the following conclusions: 1) Sinomena Zeng et al. and Yichangomena Zeng et al. are treated as junior synonyms of Eostropheodonta Bancroft; 2) Hubeinomena Zeng et al. is regarded as a juvenile form of Coolinia Bancroft; 3) Paramirorthis Zeng et al. is considered an immature growth stage of Mirorthis Zeng; 4) Shanomena Cocks and Fortey is treated as a junior synonym of Paromalomena Rong; 5) Scenidium? medlicotti Reed is redesignated as the type species of Kinnella Bergström to replace Hirnantia? kielanae Temple. The distribution of the Hirnantia Fauna in Myanmar, Thailand and western Yunnan of the Sibumasu Terrane highlights the distinctive nature of the Mandalay fauna, and the analyses of the Hirnantia Fauna and others of Sibumasu shows that Sibumasu was not located far from the South China and Lhasa palaeoplates during late Middle and Late Ordovician.     

A low-diversity,silicified Hirnantia Fauna from Butuo,southwestern Sichuan,on the western margin of Yangtze Platform

The latest Ordovician Hirnantia Fauna is distributed worldwide including South China, but with very few records from the western margin of the Yangtze Platform. A silicified Hirnantia Fauna from the uppermost Tiezufeike Formation in the Laoga section, Butuo County, southwestern Sichuan, enriches the fauna’s record and warrants the recognition of the upper part of the Tiezufeike Formation as the Kuanyinchiao Bed. Etching bulk limestone samples from this level with acetic acid yielded 275 specimens of Dalmanella testudinaria, Plectothyrella crassicosta, and Hindella crassa. The population dynamics are analyzed for three successive populations in the section. Size-frequency histograms and survivorship curves indicate gradual improvement of the palaeoenvironments during the end of the Late Ordovician glaciation. The low-diversity brachiopod assemblages recognized in our study are assigned to the Dalmanella-Plectothyrella Community (DP Community) according to their dominant species. It is comparable with two other examples of DP communities, one from the neighboring Guizhou Province of Southwest China, and the other from the Baltica Region.     

Late Darriwilian–early Katian (Ordovician) ostracods of northeastern Yunnan,Southwest China

Ordovician silicified ostracods of the western Yangtze Platform, South China, are documented for the first time from the Huadan Formation of Qiaojia County, northeastern Yunnan Province, Southwest China. Twenty-one species of 17 genera are identified with new taxa including Vendona spinifera n. sp. and Nikitinella songliangensis n. sp. These two species, together with Medianella gigantea are the most common constituents of the fauna. Faunal comparison shows close connections between South China, Baltica, the Argentine Precordillera, Kazakhstan, Tarim, and Sibumasu during this time interval. This fauna is considered to be related to a nearshore carbonate environment, and biostratigraphical analysis of it supports a late Darriwilian to early Katian age for the Huadan Formation.     

Ecological disruption close to the Ordovician‐Silurian boundary

Following a long period of stability there was substantial ecological disruption in the late Ordovician, accompanied by widespread extinction. The ecological upheaval was spread over a period of time (1 My?) and varied in intensity between different groups.

The greatest change in brachiopod faunas occurred in the tropics, where there were widespread extinctions. In temperate regions the community structure, although disrupted, was not destroyed. The deep shelf associations (Benthic Assemblages 4–5) of the Rawtheyan persisted in an attenuated form into the Hirnantian when they were supplemented by genera of the ”Hirnantia fauna” and then became strongly re‐established in the Llandovery.

Trilobites, in contrast to brachiopods, suffered a worldwide extinction in the late Ashgill. Approximately 40% of trilobite genera became extinct at, or near, the Rawtheyan‐Hirnantian boundary.

Analysis of the distribution of Rawtheyan trilobite and brachiopod genera supports a correlation between geographical distribution and survival. Global range charts for brachiopod and trilobite genera, incorporating ecological and palaeogeographical data, have been compiled for this study.     

Biostratigraphic control of the latest-Ordovician glaciogenic unconformity in Alnif (Eastern Anti-Atlas, Morocco), based on brachiopods

In the Alnif region of the Eastern Atlas (Morocco), seven fossiliferous horizons within the Lower-Ktaoua and Upper-Tiouririne formations (Ktaoua Group), as well as in the glaciomarine microconglomeratic shales of the Upper Formation of the Second-Bani Group have yielded biostratigraphically significant brachiopods and other taxa, such as trilobites and echinoderms. Several brachiopod species with short stratigraphic ranges, well-known in south-western Europe, allow a precise chronostratigraphic control of a succession that displays important lateral lithological and facies changes, when compared with the type sections in the Central Anti-Atlas. They have also permitted a better consensus between the macrofauna-determined age and that based on micropaleontological analyses. For the first time, the occurrence of a Hirnantia Fauna within the microconglomeratic shales of the Upper Formation of the Second Bani Group is reported. The biostratigraphic conclusions restrict the age of the Latest Ordovician glaciation to the early Hirnantian.     

A biogeographically mixed, Middle Permian brachiopod fauna from the Baoshan Block, western Yunnan, China

A small brachiopod fauna is described from the carbonate rocks of the basal Shazipo Formation of the Baoshan Block, western Yunnan, south-west China, including significant new ventral and dorsal internal morphological features of Cryptospirifer omeishanensis Huang. This fauna is regarded as Wordian (Middle Guadalupian, Middle Permian) because of the presence of Cryptospirifer omeishanensis Huang and associated fusulinids ( Neoschwagerina craticulifera Zone). Palaeobiogeographically, the brachiopod fauna is of considerable interest because of its admixed nature characterized by typical warm-water Cathaysian elements intermingled with temperate Peri-Gondwanan taxa. This in turn is interpreted to indicate that the Baoshan Block may have been situated in an intermediate palaeogeographical position between Gondwanaland to the south and Cathaysia to the north during the Mid Permian and, as such, it probably furnished an important 'stepping stone' for the dispersal of Mid Permian eastern Tethyan marine invertebrate taxa (e.g. Cryptospirifer ) to the western Tethys.     

An upper Kungurian/lower Guadalupian (Permian) brachiopod fauna from the South Qiangtang Block in Tibet and its palaeobiogeographical implications

A brachiopod fauna including 15 species belonging to 14 genera is described from a thin carbonate succession of the Lugu Formation at the Anmu section in the northern part of the South Qiangtang Block, which is about 15 km south to the Longmu Co-Shuanghu Suture. The brachiopod fauna is late Kungurian or early Guadalupian in age in terms of the presence of the brachiopod species Vediproductus punctatiformis, “Cryptospirifer” omeishanensis, and Paraplicatifera regularis. This is also generally consistent with the age indicated by the fusulines (e.g., Cancellina, Chusenella, Monodiexodina, Nankinella, Neofusulinella, Pseudofusulina, and Sphaerulina) from this formation at the nearby Mari and Duoma A (DMA) sections. Palaeobiogeographically, the brachiopod fauna exhibits a typical Cathaysian affinity and all the species are reported commonly from the equivalents in South China and Cimmerian blocks, which strongly suggest that the South Qiangtang Block had drifted to a position in the warm-water or temperate regime close to South China.     

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.     

Early Devonian (Lochkovian) trilobites and brachiopods from the west Balkhash Region,Kazakhstan and their significance for regional terrane dynamics

A moderately diverse Early Devonian (Lochkovian) trilobite and brachiopod association has been recovered from the lower part of the volcano-sedimentary Shakshagaily Formation, exposed on the western side of Lake Balkhash in central Kazakhstan. Its discovery demonstrates the presence of Lower Devonian marine deposits in the region. The trilobite assemblage includes representatives of six genera: the dalmanitid Kasachstania alperovichi nov. sp. is the dominant taxon, with other components being Ananaspis?, Leonaspis?, Maurotarion, Trimerus (Edgillia), and Warburgella. Associated brachiopods include species of Glossoleptaena, Leptaena, Protolepstostrophia, Pseudostrophochonetes, and Resserella. Whilst a significant proportion of these taxa show distinct links with the contemporaneous trilobite and brachiopod faunas of the north Balkhash Region, the occurrence of Trimerus (Edgillia) aff. kinglakensis Gill, 1949 suggests a possible link with the Lochkovian trilobite fauna of the Australasian sector of Gondwana. The west Balkhash Region preserves an extensive record of late Silurian to Early Devonian island arc volcanism, slope-rise sedimentation and olistostrome formation, which has invariably been ignored in models favouring early amalgamation of the Kazakhstanian terranes into a single continent by the end of the Ordovician.     

Latest Ordovician graptolites from the Mandalay Region,Myanmar

The graptolite fauna of the Mandalay Region, Myanmar, is poorly known. In this paper, 13 species in the genera Avitograptus, Korenograptus, Metabolograptus, Neodiplograptus and Normalograptus are described from the uppermost Ordovician strata of the area. Two of these species, Korenograptus selectus X. Chen n. sp. and Neodiplograptus mandalayensis X. Chen n. sp., are new. On the basis of the graptolite faunas, a biostratigraphic scheme for the uppermost Ordovician (Hirnantian: Metabolograptus extraordinarius Biozone and Metabolograptus? persculptus Biozone?) strata in this area is erected.     

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.     

The latest Ordovician Hirnantia Fauna (Brachiopoda) in time and space

The diachronous temporal and spatial distribution of the Hirnantia brachiopod fauna and the complicated pattern of terminal Ordovician events are documented through biostratigraphical analysis of the Ordovician-Silurian boundary strata in S China, Sibumasu, Xizang and elsewhere. The duration of these events (longer than the half Myr derived from isotopic excursions) indicates that they were not abrupt and instantaneous. The presence of some core taxa of the Hirnantia fauna in the upper P. pacificus Biozone (known from their earliest occurrence in China) signals the start of increased water ventilation due to the invasion of cool water across the Yangtze Basin. Low- and higher-diversity Hirnantia faunas related to onshore, shallow-water and to offshore, deeper-water environments, respectively, developed first in the basal and upper N. extraordinarius-N. ojsuensis Biozone. Disappearance of most of the fauna in the early N. persculptus Biozone suggests that the glacial maximum started to decline. The presence of the Hirnantia fauna in the upper N. persculptus to the lower P. acuminatus biozones indicates the continuation of cool water environments in some places. The diachronous disappearance of deteriorating environments (earlier in later Hirnantian and finally in the early Rhuddanian) is associated with geographical heterogeneity. Occurrences of atrypids, pentamerids and spiriferids along with key elements of the Hirnantia fauna in N Guizhou provide a link between the Late Ordovician radiation and Early Silurian recovery of these major brachiopod groups.     

The Cambrian brachiopod fauna from the first-trilobite age Shuijingtuo Formation in the Three Gorges area of China

The Yangtze platform of South China offers evidence within its Ediacaran–Cambrian geological record of the Cambrian explosion and diversification events in metazoan history. To understand the explosive radiation of animals and the environments in which it took place, the basal Cambrian fauna succession of the Aijiahe section in the Three Gorges area, western Hubei Province, has been studied, revealing the earliest brachiopod fauna (Tsunyidiscus trilobite Zone) in this region, which was dominated numerically by acrotretoids. This is accompanied by abundant skeletal fossils including minute well-preserved phosphatized archaeocyath cups and an assortment of abundant sponge spicules, chancelloriids, mollusks, hyoliths, and bradoriids, retrieved by acid-etching limestone interbeds in the black shale-dominated Shuijingtuo Formation (Series 2). The brachiopods comprise two species of acrotretoids, two types of botsfordiids (Botsfordiidae gen. et sp. indet. A and B), and four species of linguloids. Of the latter, Spinobolus popovi n. gen. n. sp. is strikingly distinctive and typified by spine-like ornamentation seen for the first time in the lower Cambrian; the remaining three linguloid genera, Palaeobolus, Eoobolus, and Lingulellotreta, have a trans-paleocontinental distribution. The Three Gorges Shuijingtuo brachiopod assemblage differs from that of the upper Atdabanian Stage (Cambrian Stage 3) in Siberia and South China, but shows great similarities with those discovered in the Tsanglangpuan (equivalent to Botoman or Stage 4) Stage of eastern Yunnan Province, Siberia, and South Australia, suggesting a much more prolonged sedimentary hiatus in basalmost Shuijingtuo Formation of the Three Gorges area than previously expected. The presence of such unconformities provides a caveat to stable isotope-based correlations that involve a number of discussions of global ocean geochemical changes across the time interval that witnessed Cambrian explosion of metazoans.     

贵州安顺募役剖面晚二叠世腕足动物群落及其生态特征*

本文对贵州募役剖面长兴期腕足动物群进行了系统分类鉴定和古生态分析,发现其与前人所报道的华南同时期腕足动物群存在着显著的差别,以Spiriferellina为绝对优势属,包含少量华南长兴期的常见分子,如Fusichonetes、Araxathyris和Peltichia等。募役剖面的腕足类化石主要产自碎屑岩层位,与华南同期碎屑岩相剖面的腕足动物群落相比,该动物群中的优势分子个体较大、壳体较厚且发育较粗大的壳疹,这可能与长兴期募役剖面动物群所处的特殊栖息地环境(浅水碳酸盐岩台地与深水硅质碎屑岩盆地之间的过渡地带)有关。二叠纪末期大规模的火山作用导致水体中碎屑物质含量增加,影响腕足类的滤食效率,而募役剖面清澈、水动力弱的水体环境,为滤食性的腕足动物提供了有利的环境条件。最后,通过与华南长兴期不同沉积相区腕足动物群落(六枝剖面、稻堆山剖面、中寨剖面、新民剖面、马家山剖面和仁村坪剖面群落)的对比分析,发现募役剖面腕足动物群与毗邻的六枝剖面腕足动物群在属级组成上的相似度较高,且过渡岩相栖息地环境下的腕足动物群在生物灭绝事件前也呈现出高优势度、低均匀度的群落结构特征,指示海洋底栖生物群落已经先于二叠纪末期生物集群灭绝事件出现了早期危机信号。     

华南与缅甸奥陶纪末赫南特贝动物群中的无铰类腕足动物

在奥陶纪末生物大灭绝的两幕之间,海洋底域繁盛着赫南特贝腕足动物群（Hirnantia Fauna）。它数量丰富,分布广泛,历程短暂。以往国内外学者研究这个动物群时,常把有铰类腕足动物作为重点,而无铰类只被简单描述或列出名单,整体面貌不明。无铰类化石尽管材料有限、研究基础薄弱,但只就属的数目而言,占据了全球赫南特贝动物群总属数的近1/5,其群落、演化和环境意义不可小觑。文中专门记述上扬子区（华南古板块）观音桥层（赫南特早中期）与缅甸曼德勒地区（滇缅马苏古地体）Hwe Mawng紫色页岩段（赫南特中期）所产赫南特贝动物群的无铰类化石,计有3目、4超科[Linguloidea（舌形贝超科）、Discinoidea（平圆贝超科）、Craniopsoidea（似髑髅贝超科）和Cranioidea（髑髅贝超科）]的10属、12种,包括5个命名属种[Plectoglossa cf. davidsoni (Barrande),Schizotretinia cf. euxina (Havlí?ek), Pseudopholidops partibilis (Rong),Petrocrania cribrum (Temple),Xenocrania haimei (Reed)]和7个未定种（Trematis sp.,Paracraniops sp.,Acanthocrania sp.,Petrocrania? sp. 1,Petrocrania? sp. 2, Pseudolingula? sp. 和Orbiculoidea sp.）,其中,Pseudolingula? sp.和Orbiculoidea sp.两个未定名种因标本不佳,未正式描述。其中,以Pseudopholidops最为常见,其次是Xenocrania和Petrocrania。前两属是华南、滇缅马苏、波罗的卡、阿瓦隆尼亚、佩鲁尼卡等古板块或地体赫南特贝动物群的常见分子。它们已知限于南、北纬30°之间的低纬度地区,这可能与化石采集和研究程度有关,但更可能反映奥陶纪末全球气候仍存在分异现象。研究识别了奥陶纪晚期无铰类腕足动物7个目,可归为3个类群：灭绝目（仅神父贝目Paterinida）、消减目（多样性与丰度大幅衰减,尤其是三分贝目Trimerellida和乳孔贝目Acrotretida在Hirnantian销声匿迹）和延续目（适应能力强、忍耐阈值高,成功穿越大灾难的首幕）。这些分类单元对研究奥陶纪-志留纪交界期腕足动物群的多样性、群落生态、生物地理及宏演化等有重要意义。     

云南德宏傣族景颇族自治州竹亚科(禾本科)植物区系地理研究

云南南部和西南部是中国竹类资源最为丰富的地区 ,与其邻近地区一起构成了世界木本竹类的多样性中心。德宏正处于这个中心位置 ,竹子种类丰富 ,有 16属 5 6种及变种 ,其中中国特有种 2 1种 (云南特有 15种 ) ,占其竹亚科全部区系成分 (45种 )的 4 6 6 7%。热带亚洲分布类型占绝对优势 ,本地区的竹亚科区系与热带亚洲特别是缅甸 (有 18种共有种 )及云南南部和东南部有非常密切的联系     

First Occurrence of Platycladus from the Upper Miocene of Southwest China and Its Phytogeographic Implications

Platycladus Spach is native to Central China, but its natural occurrences are very difficult to establish. According to molecular phylogenetic data, this genus might have originated since the Oligocene, but no fossil record has been reported. Here, we describe eight foliage branches from the upper Miocene in western Yunnan, Southwest China as a new species, P. yunnanensis sp. nov., which is characterized by foliage branches spread in flattened sprays, and leaves decussate, imbricate, scale-like and dimorphic. The leaves are amphistomatic, and the stomata are elliptical or oblong, haplocheilic, and monocyclic type. Based on a detailed comparison with the extant genera of Cupressaceae sensu lato, our fossils are classified into the genus Platycladus. The occurrence of P. yunnanensis sp. nov. indicates that this genus had a more southernly natural distribution in the late Miocene than at present. Molecular phylogeny and fossil records support a pre-Oligocene common ancestor for the genera Platycladus, Microbiota and Calocedrus. The separation of the three taxa was most likely caused by the arid belt across Central China during the Oligocene. In addition, the cooling down of the global temperature and the strengthening of Asian monsoon since the Miocene will further promote the migration of these genera.     

越南苦苣苔科石蝴蝶属植物随记暨国家级分布新记录

石蝴蝶属(Petrocosmea Oliv.)植物的现代分布与分化中心位于中国的西南山地以及周边国家和地区,如印度(阿萨姆邦)、缅甸、泰国、越南等也有分布。近年来,中国和中南半岛上各个国家和地区的新分类群和国家级新记录层出不穷,进一步加深了研究人员对中国和中南半岛植物区系以及该属植物在上述区域的多样性的理解。例如,滇黔石蝴蝶[Petrocosmea martinii (H. Léveillé)H. Léveillé]和丝毛石蝴蝶(P. sericea C.Y. Wu ex H. W. Li)曾被认为是中国的特有种,前者分布于贵州、广西北部和云南东南部,而后者仅见于云南东南部。该文报道了这两种石蝴蝶属植物的越南国家级分布新记录。这两个新记录种的发现,使得越南分布的石蝴蝶属植物种的数量上升到了5 种,这些新记录种的报道或暗示未来在与中国西南山地邻近的地区还将会有更多该属的新分类群或新记录有待发现。该文还同时提供了这两个国家级分布新记录物种的描述、图示、濒危等级现状评价以及目前所有越南已知分布的石蝴蝶属植物的检索表以备检索和核对。