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.     

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.     

Foliomena Fauna (Brachiopoda) from the Upper Ordovician of Sardinia

The late Ordovician brachiopod assemblage from Sardinia is one of the youngest members of the deep-water Foliomena fauna and is characterized by the following core taxa: Christiania , Cyclospira , Dedzetina and Foliomena . The fauna also contains Epitomyonia , Leangella , Glyptorthis and Skenidioides , which are more typical of shallower-water environments during the late Ordovician but occupied deeper-water niches during the Silurian following the termination of the Foliomena fauna. The suprafamilial placement of the family Chrustenoporidae is discussed and the new species Dedzetina serpaglii and Leangella ( Leangella ) fecunda are established. In common with many mid-Ashgill Foliomena faunas the Sardinian assemblage shows significant differences from other faunal developments of this type, reflecting its geographical position and shallower water conditions than those of the classic early Ashgill Foliomena faunas. The brachiopods occur with abundant trilobites belonging to a variant of the cyclopygid fauna. The faunas developed on part of a complex of microcontinents derived from peri-Gondwana during the Ordovician.     

A latest Ordovician Hirnantia brachiopod fauna from western Yunnan,Southwest China and its paleobiogeographic significance

A new, high-diversity, latest Ordovician brachiopod fauna of nearly 800 brachiopod specimens was collected from the Wanyaoshu Formation (Hirnantian) in the Shaodihe section, Mangshi City, western Yunnan, Southwest China. Altogether 22 genera and two undetermined taxa were identified; dominant are Aegiromena, Anisopleurella, Fardenia, Dalmanella, Hirnantia and Hindella, less common, Paromalomena, Leptaena, Eostropheodonta, Cliftonia, Kinnella, Templeella and Plectothyrella, together with some rare Petrocrania, Xenocrania, Pseudopholidops, Palaeoleptostrophia, Skenidioides, Giraldibella, Draborthis, Dolerorthis and Toxorthis. This is one of the most diverse typical Hirnantia faunas, associated with the Kosov Province. The paleobiogeographic relationships between western Yunnan (Southwest China), Myanmar, Yichang (Central China), Tibet (Southwest China) and Kazakhstan are clarified using Network Analysis and NMDS. The fauna studied is most similar to that of Myanmar; both resided on the Sibumasu terrane. However, the recalculated network diagram, when including the data of Hirnantia fauna from the Prague Basin, indicates that the latter is more closely linked to that of western Yunnan, a testament to the very weak brachiopod provincialism during the Hirnantian, mainly due to the influence of dominant cosmopolitan taxa. Some species of the fauna display significant population variation. Two of the dominant taxa, Aegiromena and Anisopleurella are systematically described, whereas the other two common taxa, Fardenia and Hirnantia are measured and their outlines and internal structures analyzed.     

Early–Mid Ordovician Yangtzeella (Syntrophiidina, Brachiopoda) and its evolutionary significance

Abstract:  Re-examination of newly collected topotype material confirms that the type species of Yangtzeella , Y .  poloi , a widespread Early and Mid Ordovician syntrophiidine brachiopod in South China, has a finely costellate shell rather than being smooth as previously thought. Thus, the subgenus Yangtzeella ( Vadimella ) Nikitina et   al ., established on the basis of fine costellae, is invalidated. Among 15 species of Yangtzeella , five species are recognized as valid based on multivariate analyses: Y .  poloi , Y .  unsulcata , Y .  songziensis , Y .  kueiyangensis and Y .  igori , among which the type species was the oldest known. Six are synonymized: Y .  septata , Y .  reticulata , Y .  lensiformis , Y .  depressa , Y .  yichangensis and Y .  minuta . Four are rejected from Yangtzeella : Y .  extensa , Y .  similior , Y .  yohi and Y .  poloi var. minor . Regional biostratigraphy indicates that Yangtzeella first appeared in a relatively deep-water setting on the Lower Yangtze Platform (South China palaeoplate) during late Tremadoc time ( Scolopodus warendensis conodont biozone) and then expanded to the deeper Jiangnan Slope as well as to the shallower Upper Yangtze Platform. The genus experienced two episodes of heightened abundance and diversity on the Upper Yangtze Platform during late Dapingian and mid Darriwilian times, respectively. Outside South China, Yangtzeella occurs sporadically in a few microplates or terranes, such as Tarim, Chu-Ili (southern Kazakhstan) and Taurides (southern Turkey) during the Dapingian and Darriwilian. Worldwide, Yangtzeella became extinct by the end of the Darriwilian.     

The earliest known strophomenoids (Brachiopoda) from early Middle Ordovician rocks Of South China

The upper Daguanshan Formation (middle Expansograptus hirundo graptolite biozone, Dapingian, early Middle Ordovician) of the Shuanghe area, Changning County, southern Sichuan Province, contains three new genera and species of strophomenoids: Ochyromena plana, Shuangheella elongata, and Primotimena globula, which are attributed to the Strophomenidae, Rafinesquinidae and Glyptomenidae respectively. These are the earliest known strophomenoids from the South China palaeoplate, and also the oldest rafinesquinid and glyptomenid brachiopods worldwide. Global review of the superfamily Strophomenoidea of Middle Ordovician age suggests that the first diversity peak at the species level occurred in late Darriwilian (Llanvirn) time, mainly as a result of the rapid diversification of the family Strophomenidae. The first appearance datum (FAD) of strophomenoids and their possible westward dispersal were from North China (latest Floian) and/or South China (early Dapingian), through the Chu‐Ili terrane of Kazakhstan, Iran, and Baltica (early Darriwilian), to Avalonia and Laurentia (late Darriwilian). This points to the existence of early diversification hotspots of the strophomenoid superfamily in the North and South China palaeoplates during the early Middle Ordovician in generally shallow water (corresponding to BA2) environments. The high degree of similarity in the external morphology and ventral interior of the three new genera indicates that the early diversification of strophomenoids began with differentiation of the cardinalia, especially in the configuration of the bilobed cardinal process, a key evolutionary novelty for the strophomenoids.     

The oldest known occurrence of the Foliomena fauna in the uppermost Darriwilian (Middle Ordovician) of South China

The oldest Foliomena fauna was, until now, known from the middle–upper Miaopo Formation (Nemagraptus gracilis Biozone, lower Sandbian, basal Upper Ordovician) of South China. In this study, the oldest record of the fauna is set back to the latest Darriwilian (upper Hustedograptus teretiusculus Biozone), represented by Foliomena jielingensis and some typical constituents of the Foliomena fauna from the basal Miaopo Formation at Jieling, northern Yichang, western Hubei Province, central China. The Miaopo Formation is characterized by its organic‐rich dark‐grey shale facies, unique in its localized distribution on the Yangtze Platform, and distinguished by its rich and diverse benthic and graptolitic faunas. This suggests an origin of the Foliomena fauna in periodically oxygen‐starved local depressions on the Yangtze Platform during the Middle–Late Ordovician transition.     

中国近海海牛亚目(软体动物)的区系分析

中国近海共有海牛亚目(Doridacea)软体动物136种,隶属于23科49属,主要分布在浙江以南的热带、亚热带海区,有些种类向北可以分布到黄、渤海,少数种类仅分布于黄、渤海,属于印度-西太平洋区的中国-日本亚区。     

Late Ordovician extinction of North American and British crinoids

After reaching a diversity peak in the Caradocian, North American Ordovician crinoids underwent a gradual decline to a nadir in the early Ashgillian (Maysvillian). This interval, recording extinction of the Cleiocrinidae, Merocrinidae, Ottawacrinidae, Hybocystitidae, and several lineages of camerate crinoids, was apparently caused by major environmental shifts in seas of eastern North America resulting from a westward-prograding wedge of terrigenous clastics derived from the Taconic Highlands, possibly coupled with a marine transgression in the Maysvillian that allowed colder water slope biofacies to invade the craton. Crinoids suffered a major episode of extinction in the late Ashgillian (late Richmondian/Rawtheyan). This event, preceding the end of the Ordovician by at least one stage or 2 to 4 million years, resulted in extinction of 12 families of crinoids including the Xenocrinidae, Tanaocrinidae, Reteocrinidae, Archaeocrinidae, Anthracocrinidae, Cincinnaticrinidae, Iocrinidae, Anomalocrinidae, Carabocrinidae, Cupulocrinidae, Porocrinidae, and Hybocrinidae. Glacio-eustatic lowering of sea level may have triggered this crisis by partially draining the North American craton, resulting in changes in oceanic circulation, salinity, and temperature. Latest Ordovician (Hirnantian) carbonates of the North American mid-continent region contain pelmatozoan assemblages from which Silurian crinoids radiated. These taxa were largely unaffected by a minor extinction event at the Ordovician/Silurian boundary.     

The influence of denitrifying seawater on graptolite extinction and diversification during the Hirnantian (latest Ordovician) mass extinction event

A continuous trench exposure within the uppermost type Vinini Formation at Vinini Creek, Roberts Mountains, Nevada, provides an unparalleled opportunity to examine the fate of graptolites, prominent Paleozoic zooplankton, during most of the Hirnantian mass extinction event. On the basis of a detailed biostratigraphic and sedimentological dataset, the relatively complete extinction record is examined in the context of ecological constraints, and it is found to reflect an ecological collapse driven by glacio-eustatic sea-level fall and associated changes in oceanic circulation. Diverse graptolite populations of the Dicranograptidae-Diplograptidae-Orthograptidae (DDO) fauna, which flourished in denitrifying waters within the oceanic oxygen-minimum zone (OMZ) during sea-level highstand, largely vanished with the loss of these conditions during glacio-eustatic sea-level fall. However, populations of one clade, the normalograptids, which inhabited the oxygenated waters of the photic zone, not only survived but diversified. These survivors gave rise to rapid recolonization and diversification with re-establishment of the oxygen-minimum and denitrifying conditions during post-Hirnantian sea-level rise. This ecological model also applies globally to other well-documented coeval stratigraphic intervals, representing both oceanic and platform sea settings.     

The earliest atrypides and athyridides (Brachiopoda) from the Ordovician of Kazakhstan

The middle Ordovician brachiopod faunas of Kazakhstan provide one of the most complete records of the evolution and radiation of some of the oldest known spire-bearing brachiopods. By contrast with North American faunas, Kazakhstanian atrypide taxa mostly belong to the suborders Atrypidina and Lissatrypidina, whereas the suborder Anazygidina is completely absent. Kazakhstanian species referred previously to ZygospiraKuzgunia are reassigned to Sulcatospira, which appeared in the Caradoc Diplograptus multidensClimacograptus clingani biozones (Sulcatospira? praecursor and Sulcatospira prima sp. nov.). Primitive, and possibly the oldest known Athyridida also appeared in Kazakhstan sometime during the Caradoc (Kellerella misiusi sp. nov.) and became widespread in brachiopod assemblages developed in carbonate mud mounds. Phylogenetic analysis suggests the early divergence of the Anazygidina, Atrypidina and Athyridida, which probably evolved independently from various primitive smooth Lissatrypidina. The new atrypide subfamily Pectenospirinae and two new atrypide genera (Rozmanospira gen. nov. and Pectenospira gen. nov. with P. pectenata sp. nov. as type species) are erected.     

Early Tremadocian (Ordovician) graptolite biostratigraphy and correlation

A revision of the early Tremadocian graptolites of the genus Rhabdinopora shows that Rhabdinopora proparabola from the Xiaoyangqiao section, Dayangcha, North China is the oldest species of the genus. Early Tremadocian species of Rhabdinopora can be differentiated easily by the construction of their nemata. The earliest species, Rhabdinopora proparabola, lacks a nema, while a multibranched nema is present in Rhabdinopora campanulatum (= Rhabdinopora parabola). The Rhabdinopora proparabola, Rhabdinopora campanulatum, Rhabdinopora flabelliformis (Anisograptus matanensis Biozone) and Rhabdinopora anglica biozones can be correlated worldwide. A short interval at the base of the Ordovician System does not contain any planktic graptolites.     

四川省蚂蚁区系研究（膜翅目：蚁科）

研究了四川（含重庆）蚂蚁的种类和分布,得知目前分布于四川的蚂蚁共有78种,分别隶属于5亚科27属,并发现1新种;古蔺立毛蚁Paratrechina gulinensis,sp.n。提供了新种的特征描述,根据地貌环境和气候的差异。将四川蚂蚁分为4个动物地理域,并讨论了四川蚂蚁区系的特点。     

Isolated Chewtonian (Lower Ordovician) graptolites from western Newfoundland

Isolated material of Chewtonian graptolites from the Cow Head Group of western Newfoundland shows some new and unexpected structural details. A high number of the two- and three-stipedPendeograptus fruticosus specimens show metasicular origin for th1¹ combined with a normal isograptid proximal development. Sigmagraptine sicular development includes a large parallel-sided prosicula and a parallel-sided metasicula. One species acquired an azygograptid-like proximal end, characterised by a loss of the initial branching, but kept the capacity to produce distal dichotomies.Maeandrograptus sinosus n. sp. andKeblograptus geminus n. sp. are described as new species.      

中国南方早更新世主要哺乳动物群层序对比和动物地理

我国南方早更新世哺乳动物群中学术界十分关注是含巨猿的"巨猿动物群"。上世纪50年代广西仅发现个别有确切层位的巨猿地点,现已扩展到鄂、桂、黔、琼和渝5省市12个地点;同时也发现了重要的人猿超科化石及文化遗存。这对探讨人类进化和古文化发展具有重要的意义。依动物群和年代学的研究,本文将早更新世哺乳动物群划分为早、中、晚三期。根据多重地层划分和对比,文中对含哺乳动物主要层位(遗址)的厘定、层序对比提出了一些不同的意见,以供讨论;同时对我国南北动物地理区系间过渡带作了必要的阐明。     

Katian (Upper Ordovician) conodonts from Wales

Middle and upper Katian conodonts were previously known in the British Isles from relatively small collections obtained from a few localities. The present study is mainly based on 17 samples containing more than 17 000 conodont elements from an approximately 14‐m‐thick succession of the Sholeshook Limestone Formation in a road cut near Whitland, South Wales, that yielded a diverse fauna of more than 40 taxa. It is dominated by representatives of Amorphognathus, Aphelognathus/Plectodina and Eocarniodus along with several coniform taxa. Representatives of Decoriconus, Istorinus and Sagittodontina are reported from the Ordovician of UK for the first time. The fauna is a typical representative of the British Province of the Atlantic Realm and includes a mixture of taxa of North American, Baltoscandic and Mediterranean affinities along with pandemic species. Based on the presence of many elements of Amorphognathus ordovicicus and some morphologically advanced specimens of Amorphognathus superbus, the Sholeshook Limestone Formation is referred to the lower A. ordovicicus Zone. Most of the unit is also coeval with Zone 2 of the Cautleyan Stage in the British regional stage classification, and stage slice Ka3 of the middle Katian Stage in the global stratigraphical classification, an age assignment consistent with data from trilobites, graptolites and chitinozoans. The unusually large collection of M elements of Amorphognathus provides insight into the complex morphological variation in this element of some Katian species of this genus. The Sholeshook conodont fauna is similar to those of the Crûg and Birdshill limestones, but differs in several respects from the slightly older ones from the Caradocian type area in the Welsh Borderland. Although having some species in common, the Sholeshook conodont fauna clearly differs from coeval Baltoscandic faunas and is even more different in composition compared with equivalent North American Midcontinent faunas.     

Reef reconstruction after extinction events of the latest ordovician in the Yangtze platform,South China

Summary Early Silurian reef reconstruction on the Yangtze Platform, in the northern part of the South China Block, is preceded by a combination of regional and global processes. During most of Ashgill time (Late Ordovician), the area was dominated by Wufeng Formation deep water graptolitic black shales. Reefs largely disappeard in the middle of the Ashgill Stage, from the northwestern margin of Cathaysian Land (southeastern South China Block), in advance of the Late Ordovician glaciation and mass extinction, due to regional sea-level changes and regional uplift, unrelated to the mass extinction itselt. Late Ordovician microbial mudmound occurrence is also found in the western margin of the Yangtze Platform, its age corresponding to theDicellograptus complexus graptolite biozone of pre-extinction time. On the Yangtze Platform, a thin, non-reef-bearing carbonate, the Kuanyinchiao Formation (=Nancheng Formation in some sites), thickness generally no more than 1m, occurs near several landmasses as a result of Hirnantian regression. Reappearance of the earliest Silurian carbonates consisting of rare skeletal lenses in the upper part of Lungmachi Formation, are correlated to theacensus graptolite biozone, early Rhuddanian of Shiqian, northeastern Guizhou, near Qianzhong Land. Carbonate sediments gradually developed into beds rich in brachiopods and crinoids in the lower part of Xiangshuyuan Formation, middle Rhuddanian. In the middle part of Xiangshuyan Formation, biostromes, containing abundant and high diversity benthic faunas such as corals, crinoids and brachiopods, show beginnings of reconstruction of reef facies. Substantial reef recovery occurred in the upper part of Xiangshuyuan Formation, lower Aeronian, as small patch reefs and biostromes. During the late Aeronian, carbonate sediments, especially reefs and reef-related facies, expanded on the upper Yangtze Platform, and radiation of reefs occurred in Ningqiang Formation, upper Telychian. The long period of reef recovery, taking several million years, remains difficult to explain, because redistribution of any refugia faunas would be expected to take place soon after the extinction. Reefs and reef-related facies subsequently declined after Telychian time due to regional uplift of the major portion of the Yangtze Platform. Carbonate facies are therefore uncommon in South China during the rest of Silurian time.     

The Upper Tremadocian (Ordovician) graptolite Bryograptus: taxonomy, biostratigraphy and biogeography

Abstract:  The taxonomy, biostratigraphical and palaeogeographical distribution of the Lower Ordovician graptolite genus Bryograptus is evaluated. Bryograptus is recognized as a distinct triradiate anisograptid with a multiramous, pendent rhabdosome. The species of the genus Bryograptus can be interpreted as shallow water faunal elements with a strongly limited biogeographical distribution to the Atlantic Faunal Realm. Bryograptus is restricted to a narrow interval in the Upper Tremadocian, the Bryograptus Biozone of Scandinavia and South America (Argentina), making it a taxon with a high potential for precise biostratigraphical correlations. The proximal end development can be used to differentiate the genus Bryograptus from other pendent multiramous graptoloid genera with a homoplastic rhabdosome development. Characteristics of the proximal end development and structure easily differentiate these genera in relief specimens, but not in flattened material.     

Victims of the Early Toarcian anoxic event: the radiation and extinction of Jurassic Koninckinidae (Brachiopoda)

The significant mass extinction attributed to the Early Toarcian anoxic event had a severe impact on the phylum Brachiopoda. Beyond a serious decrease of species diversity, the extinction of the orders Spiriferidina and Athyridida is connected with this episode. The order Athyridida was represented by the family Koninckinidae in the Early Jurassic. The stratigraphical and geographical distribution of the three Early Jurassic koninckinid genera ( Koninckella, Koninckodonta, Amphiclinodonta ) shows a definite radiative pattern. The number of their nominal species increased from 2 to 17 from the Sinemurian to Early Toarcian; in the same time interval, their area increased from the Alpine region to the whole Mediterranean and the NW-European domains. This radiative evolution can be explained as the result of different factors: (1) morphological adaptation to muddy bottoms, (2) fundamental changes in the current pattern in the Tethys/Laurasian Seaway, and, possibly, (3) utilization of methane-based chemosynthesis as alternative food source. The radiation of koninckinids, leading from the cryptic habitats of the Tethyan rocky floors to the extensive muddy bottoms of the open European shelves, was abruptly terminated by the anoxic event in the Early Toarcian Falciferum Zone. The main causes of the extinction might be: (1) the excessive warming of Tethyan deep waters by thermohaline circulation, (2) the anoxic event, which was not survived by the spire-bearers, handicapped by their stiff, calcareous spiralia. Brachiopoda, Early Jurassic, Europe, extinction, Koninckinidae, radiation, Tethys.