Testing reduced evolutionary rates during the Late Palaeozoic Ice Age using the crinoid fossil record

In 2003, Stanley & Powell reported depressed rates of origination and extinction in marine invertebrates during the Late Palaeozoic Ice Age (LPIA). Using a database of crinoid genera, rates of origination, extinction and genus duration were calculated at the stage level from the Early Devonian to the Late Permian. This 165 m.y. time span includes non‐glacial intervals before and after the LPIA, which spanned the Serpukhovian to Sakmarian, providing background rates for comparison. Data generated on crinoid evolutionary rates during the Middle to Late Palaeozoic were analysed and compared to Stanley & Powell's data to determine whether crinoid evolutionary patterns support their findings or suggest an alternative hypothesis. Rates of origination and extinction in all crinoid clades were reduced during the LPIA compared to the combined background intervals before and after the LPIA. However, crinoid diversity was higher during the LPIA than the surrounding time intervals. The difference in diversity trends between crinoids and other marine invertebrates is due to the advanced cladids clade. Unstable, fluctuating environmental conditions during the LPIA may have created habitats suitable for opportunistic crinoid genera that reduced both the probability of origination and extinction. The increased diversity of the advanced cladids is likely due to their unique adaptation of muscular arm articulations, which allowed them to thrive in marine settings with increased siliciclastic influx brought on by the Alleghenian orogeny. Despite the advanced cladids’ departure from the expected diversity count, the results of analyses performed on the updated crinoid database provide independent confirmation of Stanley & Powell's original hypothesis of depressed evolutionary rates in marine invertebrates during the LPIA.     

Collicrinus excavatus (Hall, 1861), generic affinity for a Lower Mississippian crinoid (Phylum Echinodermata)

Platycrinites was long a “catch-all” genus that was loosely defined in North America. Ausich and Kammer (2009) rectified the inconsistencies of generic diagnoses to define the Platycrinitidae better; however, due to preservation, many species were not known from specimens with sufficient characters to be placed confidently into a genus and, thus, were designated Platycrinites sensu lato. Collicrinus excavates (n. comb.) was one such species. A specimen with a complete theca and partial arms preserved was discovered in southwestern Missouri in the Burlington–Keokuk Limestone. With the discovery of this specimen, C. excavatus has been fully described and placed in the correct genus. Further, the associated fauna of new C. excavatus specimens, reported here, helps to evaluate the stratigraphic position of this species, which is now considered to be only from Burlington Pelmatozoan Association I and II, roughly equivalent to the Dolbee Creek and Haight Creek Members of the Burlington Limestone.     

Mississippian crinoid biodiversity,biogeography and macroevolution

Abstract: The biodiversity and biogeography of 217 genera of Mississippian crinoids from North America and the British Isles shed light on the macroevolutionary turnover between the Middle Palaeozoic and Late Palaeozoic Crinoid Evolutionary Faunas. This turnover resulted from steady differential extinction among clades during the middle Mississippian after crinoids reached their Phanerozoic peak of generic richness during the early Mississippian. This peak richness was primarily a function of Mississippian originations rather than Devonian–holdover taxa. North America had 100 per cent higher generic richness than the British Isles, but rarefaction analysis adjusts the difference to only 37 per cent higher. Rarefaction demonstrated that North America had increased biodiversity, compared to the British Isles, almost entirely among monobathrid camerates, disparids and primitive cladids. In contrast, diplobathrid camerates, advanced cladids and flexibles had the same generic biodiversity between regions, when compared using rarefaction. The early Mississippian radiation resulted from two primary causes: (1) the expansion of Tournaisian carbonate ramps following the Frasnian mass extinction of reef faunas and (2) the predatory release in the Tournaisian following the end‐Famennian Hangenberg extinction of durophagous fishes. A majority of crinoid genera from the British Isles are cosmopolitan. When combined with rarefaction analysis and evidence for more first occurrences in North America, this suggests higher origination rates in North America, especially when carbonate ramps were widespread. With the gradual reduction in the area of carbonate ramps from the early to late Mississippian, in conjunction with the radiation of new durophagous fishes, camerate crinoids in particular experienced continuous background extinction, without replacement, beginning during the earliest Viséan (late Osagean). By middle Viséan time (late Meramecian) advanced cladids were dominant in all settings. This resulted in the transition from the Middle Palaeozoic to the Late Palaeozoic Crinoid Macroevolutionary Fauna.     

New data on the distribution of the Upper Carboniferous (Pennsylvanian) flexible crinoid Cibolocrinus (Crinoidea,Flexibilia)

Two new species of flexible crinoids of the genus Cibolocrinus (Crinoidea, Flexibilia) from the Upper Carboniferous of the Moscow Region are described: C. kutasovi (Moscovian Stage, Podolskian Regional Substage) and C. gerassimovi (Gzhelian Stage, Dobryatinian Regional Substage). The first species, C. kutasovi, belongs to a group with low cone shaped cups that first appeared in the Moscow Basin and then spread to the Midcontinent of North America. In one specimen of C. gerassimovi the smallest infrabasal plate is situated in the A ray, which is not typical for flexible crinoids. A poorly preserved crown of Cibolocrinus sp. from the Upper Carboniferous (Gzhelian Stage) of the Darvaz Ridge (western Pamir, central Tajikistan) is also described. The described specimens of Cibolocrinus from the Moscow Region and the Darvaz Ridge are the first reliable specimens of this genus described from Europe and Asia.     

Aspects of crinoid palaeontology of the North Esk Inlier,Scotland (Silurian,Llandovery, Telychian)

Abstract: The described fauna of well‐preserved Llandovery (Telychian) echinoderms from the North Esk Inlier, including six crinoids, one echinoid and seven starfish species, is mainly allochthonous. Most of these taxa are known only from starfish beds, channel fill deposits probably representing submarine mass flows and preserving a biota probably derived from elsewhere, presumably shallower water. Only one crinoid species, Pisocrinus cf. campana Miller, is recognized as a common fossil away from the starfish beds and is a biostratigraphic marker for the base of the Wether Law Linn Formation, forming part of the Skenidioides‐Cyrtia Association. Crinoid columnals preserved perpendicular to bedding (that is, in putative life position) in Lamont’s bivalve bed, Deerhope Formation, are tentatively interpreted as being in situ by comparison with a similar occurrence in the Silurian of Arisaig, Nova Scotia. Two new species of crinoid are described, the cladid Dendrocrinus? sp. and the columnal morphospecies Pentagonocyclicus (col.) lamonti sp. nov.     

Tube foot preservation in the Devonian crinoid Codiacrinus from the Lower Devonian Hunsrück Slate,Germany

Fossilized tube feet are described on Codiacrinus schultzei Follmann from the Lower Devonian Hunsrück Slate of Germany. This is the first definitive proof of tube feet on any fossil crinoid. Three lightly pyritized, flattened tube feet are preserved in a single interray of this cladid crinoid. The tube feet were at least 7 mm long. Their preservation is very similar to the tube feet reported previously from a Hunsrück ophiuroid, except that the Codiacrinus tube feet have small papillae, similar to living crinoids.     

Radial perforations in crinoid stems from the Silurian of Gotland

Radiating intracolumnal canals are a characteristic feature of large (diameter 10 mm or more) crinoid stems from the Silurian of Gotland. They are found in nodals as well as in internodals where the columnal height exceeds one millimetre. They were formed secondarily in the median and distal portions of crinoid stems with pseudocirriferous holdfasts. Intercolumnal canals are found in the distal parts of stems with true cirri regardless of the size of the stem. It is suggested that these canals played an important role in crinoid physiology. The crinoids are believed to have sustained a large proportion of their tissues through cutaneous digestion and uptake of dissolved substances from the surrounding sea water. The intra- and intercolumnal canals increased the surface of the axial canal in relation to volume. They provided a connection between the axial canal and the surrounding sea water, thus facilitating nutrient transport to the tissues.     

The validity of the Middle Devonian camerate crinoid species Hexacrinites hieroglyphicus (Goldfuss, 1839) and H. marginatus (Schultze, 1866)

The camerate crinoids Hexacrinites hieroglyphicus (Goldfuss, 1839) and H. marginatus (Schultze, 1866) are valid species known from the Lower Givetian of the synclines of Gerolstein and Hillesheim, Eifel Hills (westernmost Germany). As shown by the refigured holotypes and new material, they are mostly separated by the expressed ornamentation of radials and basals. The radials of H. hieroglyphicus are characterized by prominent, variously formed bulges, stretching preferably in a radiate pattern across most of the plates. They do not reach the low plate boundaries. Vice versa, H. marginatus shows expressed depressions in the central part of the plates, but raised plate margins. The first known crowns of H. marginatus are described; one of them is proposed as neotype for the lost original.     

Diversity dynamics of post‐Palaeozoic crinoids – in quest of the factors affecting crinoid macroevolution

Following the end‐Permian biotic crisis which led to the near extinction of crinoids, this echinoderm class rebounded rapidly during the Mesozoic, resulting in forms with important morphological and behavioural novelties. However, quantitative patterns of crinoid diversity during the Mesozoic remain largely unexplored. Here, we report results of analyses of the evolutionary dynamics of post‐Palaeozoic crinoid genera spanning a time interval between 250 and 70 Myr. We show that crinoids reached their Mesozoic peak of genus‐level richness during the Late Jurassic. We also document a major reorganization of different ecological crinoid groups in the Mesozoic. More specifically, the diversity of sessile forms generally increased towards the mid‐Mesozoic but decreased significantly starting in the Cretaceous, whereas the number of motile crinoid genera increased linearly during the Mesozoic. The possible role of biotic and abiotic factors in crinoid evolution is discussed.     

Belling the Chinese Dragon at Sea: Western Theories and Asian Realities

Against the backdrop of projections of either conflict escalation or legal settlement by Western international relations theories, this article provides a more nuanced and subtle balance between Western theories and Asian realities.     

New crinoids from the Baltic region (Estonia): fossil tip‐dating phylogenetics constrains the origin and Ordovician–Silurian diversification of the Flexibilia (Echinodermata)

This study documents previously unknown taxonomic and morphological diversity among early Palaeozoic crinoids. Based on highly complete, well preserved crown material, we describe two new genera from the Ordovician and Silurian of the Baltic region (Estonia) that provide insight into two major features of the geological history of crinoids: the early evolution of the flexible clade during the Great Ordovician Biodiversification Event (GOBE), and their diversification history surrounding the end‐Ordovician mass extinction. The unexpected occurrence of a highly derived sagenocrinid, Tintinnabulicrinus estoniensis gen. et. sp. nov., from Upper Ordovician (lower Katian) rocks of the Baltic palaeocontinent provides high‐resolution temporal, taxonomic and palaeobiogeographical constraints on the origin and early evolution of the Flexibilia. The Silurian (lower Rhuddanian, Llandovery) Paerticrinus arvosus gen. et sp. nov. is the oldest known Silurian crinoid from Baltica and thus provides the earliest Baltic record of crinoids following the aftermath of the end‐Ordovician mass extinction. A Bayesian ‘fossil tip‐dating’ analysis implementing the fossilized birth–death process and a relaxed morphological clock model suggests that flexibles evolved c. 3 million years prior to their oldest fossil record, potentially involving an ancestor–descendant relationship (via ‘budding’ cladogenesis or anagenesis) with the paraphyletic cladid Cupulocrinus. The sagenocrinid subclade rapidly diverged from ‘taxocrinid’ grade crinoids during the final stages of the GOBE, culminating in maximal diversity among Ordovician crinoid faunas on a global scale. Remarkably, diversification patterns indicate little taxonomic turnover among flexibles across the Late Ordovician mass extinction. However, the elimination of closely related clades may have helped pave the way for their subsequent Silurian diversification and increased ecological role in post‐Ordovician Palaeozoic marine communities. This study highlights the significance of studies reporting faunas from undersampled palaeogeographical regions for clade‐based phylogenetic studies and improving estimates of global biodiversity through geological time.     

中国古生代放射虫研究世纪回顾—为庆贺盛金章院士80华诞而作

20世纪中国古生代放射虫研究取得三方面成绩：1.研究地域涉及全国大部分省区。除东北三省、河北、山西、山东、海南、台湾外,古生代放射虫巳在我国19个省区发现。时代从晚寒武世至晚二叠世长兴期;2.研究领域逐步扩大,确立了三种不同的放射虫岩相类型,洋盆相区以广西钦州板城最为典型,从晚泥盆世弗拉斯期-晚二叠世长兴早期共建立18个化石带;岛孤相区以云南西部昌宁-孟连地体作代表,从中泥盆世吉维持期-晚二叠世长兴期发育12个化石带;台盆相区以苏皖地区中二叠世孤峰组放射虫研究得较好,建立3个化石带,这些放射虫化石带,与洋盆相区,岛弧相区同时代的带基本上相似和相同;3.中国4条主要蛇绿岩带都发现了放射虫硅质岩和放射虫动物群。这对蛇绿岩带的形成时代和板块碰撞时间和确定提供了有力的证据。最后,文中还提出了新世纪里我国古生代放射虫研究的方向。     

China,Vietnam, and the South China Sea: Disputes and Dispute Management

This article examines recent developments in the South China Sea; in particular, the China-Vietnam relationship. The developments are presented in the broader context of the Sino-Vietnamese approach to managing border disputes since full normalization of relations in late 1991. The challenges for China and Vietnam in managing their disputes and related tension in the South China Sea are also discussed.     

Setting Up International (Adversary) Regimes in the South China Sea: Analyzing the Obstacles from a Chinese Perspective

If the sovereignty issue can be shelved, solved, or resolved, the six claimant states of the South China Sea will have to establish a series of international regimes. The following questions need to be answered: Do all the claimants understand the concept of international regimes? Would the regimes be imposed? Would there be hegemon(s) in the South China Sea? If so, which country or multiple entities will be involved? What would be the outcome of having international regimes in the South China Sea? Can the experience elsewhere be applied to the South China Sea?     

China's Nine Dotted Lines in the South China Sea: The 2011 Exchange of Diplomatic Notes Between the Philippines and China

Important events relating to the sovereignty dispute over the Spratly Islands have arisen by fits and starts since 2009, marking the start of a new phase in the legal battle over territorial and maritime claims in the South China Sea. While the exchange of legal arguments between the parties has gradually laid bare their maritime claims, much still remains shrouded in uncertainty. Among the obscure claims wanting clarification is China's infamous nine-dotted-line map, which in 2011 elicited a response and counterresponse between the Philippines and China. This article examines the maritime and territorial claims of the Philippines and China as revealed in the recent discord over the nine-dotted-line map.     

Taxonomical notes on the family Ptilocodiidae (Anthomedusae) from the central and southern of South China Sea,with a new genus and a new species

This study reviews the Ptilocodiidae from South China Sea,including three species.The diagnoses of the family and each genus are provided.A new genus,Tregouboviopsis Guo,Xu & Huang,gen.nov.is erected,based on the monotypy,Tregouboviopsis perradialis (Xu,Huang & Du,2012) comb.nov.,which is transferred from the genus Tregoubovia.A free eumedusoid ofHydrichthella ocellata Xu,Huang & Wang,sp.nov.is described in the present paper.A key to known genera of family Ptilocodiidae is presented.All type specimens are deposited in the South China Sea Fisheries Research Institute,Chinese Academy of Fishery Science.     

Book review

H. D. Holland, The Chemical Evolution of the Atmosphere and Oceans. Princeton University Press, Princeton, N.J., 1984. 94 tables and 230 figures, xii + 582 pp., $75.00; paper $24.00.     

Coastal States in the South China Sea and Submissions on the Outer Limits of the Continental Shelf

The purpose of this article is to examine the outer continental shelf submissions made by the coastal states of the South China Sea and their potential impact on legal and political developments in the South China Sea. In accordance with the United Nations Convention on the Law of the Sea of 1982 and the guidelines of the Commission on the Limits of the Continental Shelf, coastal states are to establish the outer limits of their continental shelf where it extends beyond 200 nautical miles. Meeting this obligation is complicated in enclosed or semienclosed seas where there are maritime disputes such as in the South China Sea.     

A Code of Conduct for Human and Regional Security Around the South China Sea

In the past decade, there have been calls for a regional code of conduct for the South China Sea to ensure peace and stability in a region replete with conflicting territorial claims over offshore geographical features. This paper will distinguish between the process of codification of rules and principles in one document, on the one hand, and the process of implementing existing rules and principles, on the other hand. It will be argued that rules and principles governing conduct for human and regional security around the South China Sea already exist in many forms and that the South China Sea states are obligated by international law to implement them without waiting for their consolidation in one document.