Diversity and extinction patterns of permian brachiopoda of South China

The stratigraphical and geographical distribution of 851 brachiopod species from 216 genera and 65 families in the Permian of South China are analysed. It is revealed that the brachiopod diversity underwent two sharp falls during the Permian. The first occurred at the end of Maokouan, accompaning the widely recognised, extensive regression across the Maokouan‐Wujiapingian boundary. Fifty‐seven species of 29 genera survived this first major extinction event. The second sharp reduction of brachiopod diversity took place in the later Changhsingian, with only 17 Permian‐type brachiopod species of 12 genera straggling into the earliest Triassic. Detailed stratigraphic analysis shows that more than 90% of the Changhsingian brachiopod species disappeared at different levels in the Changhsingian before the widely perceived end‐Permian ‘mass extinction’ occurred. It is also notable that each of the step‐wise diversity reduction events was apparently heterochronous. In view of the evidence from lithologies, faunal components and geochemical analyses, the two sharp falls of Permian brachiopod diversity in South China are considered to be closely related to multiple interactions of an environmental deterioration caused by large‐scale regressions.     

浙江煤山剖面二叠纪末的小有孔虫动物群

通过对浙江长兴煤山剖面长兴阶上部和二叠系三叠系界线地层进行系统切片和研究,共鉴别出小有孔虫化石18属50种。这一丰富的有孔虫动物群,不仅进一步完善了长兴阶标准剖面的化石资料,而且也为研究二叠纪末的生物灭绝形式提供了新的依据。     

The Permian/Triassic mass extinction among brachiopods in the Northern Caucasus (northern Palaeo-Tethys): A tentative assessment

A palaeontological record of the Northern Caucasus provides new data to evaluate the influence of the Permian/Triassic mass extinction on brachiopod communities. The study region is currently located in the southwest of Russia; it laid on the northern margin of the Palaeo-Tethys Ocean during the late Paleozoic-early Mesozoic. A total of 168 genera and 36 superfamilies are known from the Changhsingian-Bajocian deposits of this region. The total diversity of brachiopods was very high in the Changhsingian (57 genera and 19 superfamilies), but these organisms disappeared entirely at the Permian/Triassic boundary. Three genera and three superfamilies existed in the Induan, but brachiopods perished again in the Olenekian. A recovery began in the Anisian, but the Changhsingian diversity was never reached again. No genera crossed the Permian/Triassic boundary, whereas 4 superfamilies became able to do this. Ambocoelioidea was a “dead clade walking” and went extinct in the Induan, whereas three other superfamilies (Dialasmatoidea, Rhynchonelloidea, and Wellerelloidea) were more successful. Survivors included no less than 20% of genera during the entire Induan-Bajocian time interval. The Changhsingian-Anisian interval is dominated by just one type of facies in the studied region, which may explain differences in the post-extinction diversity patterns between the Northern Caucasus, South China, and other parts of the World. Use of the alternative Triassic time scale does not change the absolute duration of the post-extinction stress (5.1 myr), whereas it highlights higher rates for the Anisian recovery (appearance of 0.9 superfamilies and 2.8 genera per myr).     

Extinction process and patterns of Middle Permian Fusulinaceans in southwest China

Study of the extinction process of 179 fusulinacean species belonging to 34 genera in southwest China demonstrates that diversity changes in the small (shell length ≤6  mm) and large (shell length >6  mm) species groups were very similar throughout the 'Maokouan' (∼Guadalupian). However, significant differences in the timing of extinction pulses occur between different fusulinacean clades with different wall structures, i.e. the nankinellids, schwagerinids, verbeekinids and neoschwagerinds, and between the large and small species groups within the schwagerinids and verbeekinids. Fusulinacean diversity reveals that the Guadalupian mass extinction began in the Middle 'Maokouan' and greatly intensified in the late 'Maokouan'. With only seven species of five genera surviving into the Late Permian, the extinction of fusulinaceans in the Guadalupian mass extinction is 96% at species level and 85% at generic level in southwest China. The preferential extinction of large, morphologically complicated species and the survival of simpler, ecologically more tolerant species of the nankinellids suggest that the extinction of Guadalupian fusulinaceans was caused by falling sea level and the consequent effects related to salinity, temperature and substrate changes. It is also demonstrated that biological characters of fusulinaceans such as shell size and test structures could have certain effects on survivorship of species in the early stages of mass extinction when extinction pressure was less intensive but were ineffectual during the extinction climax.     

Discovery of the Late Changhsingian Bivalve Complex and Two Fauna Extinction Episodes in Northeastern Asia at the End of the Permian

A late Changhsingian bivalve complex including species from the genera Palaeonucula, Dacryomya, Malletia ?, Sarepta ?, Myalina, Pteria, Maitaia, and Unionites is discovered in northeastern Asia for the first time. The transition from the Permian to the Triassic in high-boreal basins has been shown to include two extinction episodes similar to those observed in the low-boreal basins and apparently evoked by trap volcanism activation in Siberia. Changes in benthic foraminifera diversity and vertical distribution of ammonoidea of the genus Otoceras in transitional Permian–Triassic deposits also are considered. Images of bivalves from the most typical taxa are presented.     

贵州紫云晒瓦群中放射虫Neoalbaillella动物群的发现

首次报道贵州紫云晚二叠世晒瓦群顶部硅质岩中Neoalbaillella动物群,计12种,5未定种,归属于12属9科4超科2亚目,其中1新属（Spongosphaeradiscus gen.nov.）,2新种（Spongosphaeradiscus chaiwaensis sp.nov.和Copicyntra shaiwaensis sp.nov.）。根据这个动物群建立Neoalbaillella ornithoformis-Spongosphaeradiscus shaiwaen-sis带,与东亚和美国西海岸长兴期的Neoalbaillella ornithoformis带对比。     

CALIBRATED DIVERSITY,TREE TOPOLOGY AND THE MOTHER OF MASS EXTINCTIONS: THE LESSON OF TEMNOSPONDYLS

Abstract: Three family‐level cladistic analyses of temnospondyl amphibians are used to evaluate the impact of taxonomic rank, tree topology, and sample size on diversity profiles, origination and extinction rates, and faunal turnover. Temnospondyls are used as a case study for investigating replacement of families across the Permo‐Triassic boundary and modality of recovery in the aftermath of the end‐Permian mass extinction. Both observed and inferred (i.e. tree topology‐dependent) values of family diversity have a negligible effect on the shape of the diversity curve. However, inferred values produce both a flattening of the curve throughout the Cisuralian and a less pronounced increase in family diversity from Tatarian through to Induan than do observed values. Diversity curves based upon counts of genera and species display a clearer distinction between peaks and troughs. We use rarefaction techniques (specifically, rarefaction of the number of genera and species within families) to evaluate the effect of sampling size on the curve of estimated family‐level diversity during five time bins (Carboniferous; Cisuralian; Guadalupian–Lopingian; Early Triassic; Middle Triassic–Cretaceous). After applying rarefaction, we note that Cisuralian and Early Triassic diversity values are closer to one another than they are when the observed number of families is used; both values are also slightly higher than the Carboniferous estimated diversity. The Guadalupian–Lopingian value is lower than raw data indicate, reflecting in part the depauperate land vertebrate diversity from the late Cisuralian to the middle Guadalupian (Olson’s gap). The time‐calibrated origination and extinction rate trajectories plot out close to one another and show a peak in the Induan, regardless of the tree used to construct them. Origination and extinction trajectories are disjunct in at least some Palaeozoic intervals, and background extinctions exert a significant role in shaping temnospondyl diversity in the lowermost Triassic. Finally, species‐, genus‐, and family trajectories consistently reveal a rapid increase in temnospondyl diversity from latest Permian to earliest Triassic as well as a decline near the end of the Cisuralian. However, during the rest of the Cisuralian family diversity increases slightly and there is no evidence for a steady decline, contrary to previous reports.     

云南地区种子植物多样性分布格局和多样性分化特点研究

利用大尺度的植物分布信息结合地理信息系统和数理统计分析,探讨了云南种子植物多样性的地理分布格局和多样性分化特点。结果表明,从科到属和种,植物多样性和区系分化强度的数据集离散程度均呈递增趋势。相对属和种而言,科的多样性的空间分异和高值区域并不明显,这可能与科、属、种所代表的不同地质历史时期有关;南部地区属和种的多样性明显高于除滇西北以外的大部分地区。种的多样性分布中心主要集中在滇西北地区、云南南部地区、哀牢山、无量山地区和昆明地区,这可能与生境异质性和热量条件有关。相对科而言,属的区系分化强度的空间分异更为明显,并与物种多样性的分布格局比较接近,暗示着植物多样性与区系分化之间的密切联系。相对科而言,属的区系分化中心高值区域更为明显。总体上,云南地区植物多样性的分布格局及其分化可能与其生境、热量条件和地质历史有关。     

广西南部二叠纪长兴期放射虫动物群

广西南宁南部柳桥区的长兴组上部硅质岩中存着精美的长兴期晚期放射虫动物群,共有24属39例,其中3个新种,其中以Latentifistularia最为常见,其次是Albaillllidae和Entactiniidae,属于二叠纪晚期的Neoalbaillella optima组合。半生生物有有孔虫、海绵骨针、腕足动物等。多数属在接近二叠纪－三叠世界线附近消失,但生物群中已出现少量三叠纪分子。     

Spatial pattern of coral diversity in Luhuitou fringing reef, Sanya, China

84 quadrats from 5 vertical transects of Luhuitou fringing reef are investigated in detail by using video-quadrat and indoor-interpretation methods. The results show that (1) the reef consists of 69 species of hermatypic corals belonging to 24 genera and 13 families which are unevenly distributed in abundance. (2) Among all the corals, Porites lutea is the most dominant species with importance value percentage up to 36.62%; Porites and Acropora are dominant genera with importance value percentages 43.85% and 22.88%, respectively. (3) There exist distinct spatial differences in coral communities. Both the coral covers and coral diversity indices on the northeastern transects are higher than those on the central and southern transects. (4) Coral communities also show remarkable zonal characteristics with less coral species occurring on reef flat than on reef slope. The importance value percentage of the sole dominant coral genus, Porites, is over 50%, while on the reef slope, the importance value percentages are 28.33% for the first dominant genus Acropora and 26.71% for the second dominant genus Porites. Our further analysis suggests that the spatial and zonal differences of coral diversity pattern are correlated with both natural environmental changes and human activities. The shallow water reef flat is frequently exposed at low tide and it receives more anthropogenic influences (including dredging and trampling) than the deep water reef slope. Thus, the coral community on the reef flat is not as well developed as that on reef slope. The relatively poor coral covers and coral diversity indices on the central and southern transects are closely related to heavy human activities around these sites such as aquaculture, fishing and coastal sewage drainage. Therefore, the impact of human activities must be taken into account in developing strategies for the protection of this coral reef.     

Spatial pattern of coral diversity in Luhuitou fringing reef, Sanya, China

84 quadrats from 5 vertical transects of Luhuitou fringing reef are investigated in detail by using video-quadrat and indoor-interpretation methods. The results show that (1) the reef consists of 69 species of hermatypic corals belonging to 24 genera and 13 families which are unevenly distributed in abundance. (2) Among all the corals, Porites lutea is the most dominant species with importance value percentage up to 36.62%; Porites and Acropora are dominant genera with importance value percentages 43.85% and 22.88%, respectively. (3) There exist distinct spatial differences in coral communities. Both the coral covers and coral diversity indices on the northeastern transects are higher than those on the central and southern transects. (4) Coral communities also show remarkable zonal characteristics with less coral species occurring on reef flat than on reef slope. The importance value percentage of the sole dominant coral genus, Porites, is over 50%, while on the reef slope, the importance value percentages are 28.33% for the first dominant genus Acropora and 26.71% for the second dominant genus Porites. Our further analysis suggests that the spatial and zonal differences of coral diversity pattern are correlated with both natural environmental changes and human activities. The shallow water reef flat is frequently exposed at low tide and it receives more anthropogenic influences (including dredging and trampling) than the deep water reef slope. Thus, the coral community on the reef flat is not as well developed as that on reef slope. The relatively poor coral covers and coral diversity indices on the central and southern transects are closely related to heavy human activities around these sites such as aquaculture, fishing and coastal sewage drainage. Therefore, the impact of human activities must be taken into account in developing strategies for the protection of this coral reef.     

An ecologically mixed brachiopod fauna from Changhsingian deep-water basin of South China: consequence of end-Permian global warming

The Late Permian Shaiwa Group of the Ziyun area of Guizhou, South China is a deep-water facies succession characterized by deep-water assemblages of pelagic radiolarians, foraminifers, bivalves, ammonoids and brachiopods. Here we report 20 brachiopod species in 18 genera from the uppermost Shaiwa Group. This brachiopod fauna is latest Changhsingian in age and dominated by productides. The palaeoecologic and taphonomic analysis reveals that the brachiopod fauna is preserved in situ. The attachment modes and substratum preference demonstrate that the Shaiwa brachiopod fauna comprises admixed elements of deep-water and shallow-water assemblages. The presence of the shallow-water brachiopods in the Shaiwa faunas indicates the involuntary settlement of shallow-water brachiopods. The stressed ecologic pressure, triggered by warming surface waters, restricted ecospace and short food sources, may have forced some shallow-water elements to move to hospitable deep-water settings and others to modify their habiting behaviours and exploit new ecospace in deep-water environments. We infer that the end-Permian global warming and subsequent transgression event may have accounted for the stressed environmental pressure in the shallow-water communities prior to the end-Permian mass extinction.     

Global diversity of mayflies (Ephemeroptera, Insecta) in freshwater

The extant global Ephemeroptera fauna is represented by over 3,000 described species in 42 families and more than 400 genera. The highest generic diversity occurs in the Neotropics, with a correspondingly high species diversity, while the Palaearctic has the lowest generic diversity, but a high species diversity. Such distribution patterns may relate to how long evolutionary processes have been carrying on in isolation in a bioregion. Over an extended period, there may be extinction of species, but evolution of more genera. Dramatic extinction events such as the K-T mass extinction have affected current mayfly diversity and distribution. Climatic history plays an important role in the rate of speciation in an area, with regions which have been climatically stable over long periods having fewer species per genus, when compared to regions subjected to climatic stresses, such as glaciation. A total of 13 families are endemic to specific bioregions, with eight among them being monospecific. Most of these have restricted distributions which may be the result of them being the relict of a previously more diverse, but presently almost completely extinct family, or may be the consequence of vicariance events, resulting from evolution due to long-term isolation. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Reef Fishes in Biodiversity Hotspots Are at Greatest Risk from Loss of Coral Species

Coral reef ecosystems are under a variety of threats from global change and anthropogenic disturbances that are reducing the number and type of coral species on reefs. Coral reefs support upwards of one third of all marine species of fish, so the loss of coral habitat may have substantial consequences to local fish diversity. We posit that the effects of habitat degradation will be most severe in coral regions with highest biodiversity of fishes due to greater specialization by fishes for particular coral habitats. Our novel approach to this important but untested hypothesis was to conduct the same field experiment at three geographic locations across the Indo-Pacific biodiversity gradient (Papua New Guinea; Great Barrier Reef, Australia; French Polynesia). Specifically, we experimentally explored whether the response of local fish communities to identical changes in diversity of habitat-providing corals was independent of the size of the regional species pool of fishes. We found that the proportional reduction (sensitivity) in fish biodiversity to loss of coral diversity was greater for regions with larger background species pools, reflecting variation in the degree of habitat specialization of fishes across the Indo-Pacific diversity gradient. This result implies that habitat-associated fish in diversity hotspots are at greater risk of local extinction to a given loss of habitat diversity compared to regions with lower species richness. This mechanism, related to the positive relationship between habitat specialization and regional biodiversity, and the elevated extinction risk this poses for biodiversity hotspots, may apply to species in other types of ecosystems.     

Reef coral diversity and global change

Regional anthropogenic processes such as pollution, dredging, and overfishing on coral reefs currently threaten the biodiversity of stony corals and other reef-associated organisms. Global climate change may interact with anthropogenic processes to create additional impacts on coral diversity in the near future. In order to predict these changes, it is necessary to understand the magnitude and causes of variation in scleractinian coral diversity throughout their 240 million year history. The fossil record documents long periods of speciation in corals, interrupted repeatedly by events of mass extinction. Some of these events relate clearly to changes in global climate. Diversity in reef corals has increased since their last period of extinction at the end of the Cretaceous (65 My bp ), and is still rising. During the last 8 million years, the fragmentation of the once pantropical Tethys Sea separated corals into two major biogeographical provinces. Periods of glaciation also have caused major changes in sea level and temperature. Accumulated evidence supports the theory that relative habitat area and changing patterns of oceanic circulation are mainly responsible for the two observed centres of recent coral diversity at the western tropical margins of the Atlantic and Pacific oceans. At predicted rates of climate change in the near future, coral reefs are likely to survive as an ecosystem. Increases in sea level may actually benefit corals and lead to regional increases in diversity if new habitat area on back reefs is opened to increased water circulation and thus coral dispersal. Rising temperature may cause higher rates of coral mortality and even local extinction in isolated, small populations such as those on oceanic islands. The effects of increases in ultraviolet radiation (UV) are largely unknown, but likely to be negative. UV may damage planktonic coral propagules in oceanic surface waters and thus decrease rates of gene flow between coral populations. This may result in increased local extinctions, again with the strongest impact on widely separated reefs with small coral populations. The largest threats to coral diversity are regional anthropogenic impacts, which may interact with global climate change to exacerbate rates of local species extinctions. Centres of high reef coral diversity coincide with human population centres in south-east Asia and the Caribbean, and thus the greatest potential for species loss lies in these geographical areas.     

Diversity of Scleractinia and Octocorallia in the mesophotic zone of the Great Barrier Reef, Australia

Mesophotic coral reefs in the Indo-West Pacific, the most diverse coral reef region on earth, are among the least documented. This study provides the first detailed investigation of the diversity of Scleractinia and Octocorallia of the mesophotic Great Barrier Reef (GBR). Specimens were collected by 100-m rock dredge tows at 47–163 m depth on 23 sites in four regions (15.3°–19.7° latitude South). Twenty-nine hard coral species from 19 families were recorded, with the greatest diversity found at <60 m depth, and no specimen was found >102 m. Many of these species are also commonly observed at shallower depths, particularly in inshore areas. Twenty-seven octocoral genera were collected, 25 of which represented azooxanthellate genera. Generic richness of octocorals was highest at depths >60 m. Sixteen of the 25 azooxanthellate genera were either absent or very rare at <18 m, and only five azooxanthellate genera were common on both shallow and mesophotic reefs. Species-area models indicated that the total diversity of hard corals on the deep mesophotic reefs sampled during this study was ~84 species while octocorals were represented by ~37 genera; however, the wide 95% confidence limits indicates that more intensive sampling effort is required to improve the accuracy of these estimates. Nonetheless, these results show that the taxonomic richness, particularly of hard corals, on mesophotic reefs may be much higher than previously thought, a finding that has implications for the comprehensive and adequate protection of the full range of biodiversity of the GBR.     

End-Permian mass extinction of calcareous algae and microproblematica from Liangfengya,South China

A new assemblage of calcareous algae and microproblematica is reported from the Changxing Formation at the Liangfengya section in Chongqing, South China. This assemblage comprises eighteen species of seven genera, including three genera of gymnocodiaceaens (Gymnocodium, Permocalculus, and Tauridium), three genera of dasycladaleans (Epimastopora, Macroporella, and Mizzia), and one genus of microproblematica (Pseudovermiporella). A new algal species, Tauridium elongatum nov. sp., is described. Quantitative analysis indicates that the last occurrences of 22% of the species (4 out of 18) fall into a 52 cm thick interval of the uppermost Changhsingian. No calcareous algae are found in the Permian-Triassic boundary (P-T boundary) beds and the overlying Feixianguan Formation. Four out of 10 species have a stratigraphic abundance greater than 10% and all vanished in the uppermost Changhsingian. An abrupt extinction for calcareous algae occurred in the Clarkina yini conodont Zone.     

Changhsingian brachiopod communities along a marine depth gradient in South China and their ecological significance in the end-Permian mass extinction

Diversity indices (dominance and evenness) and ecological spatial structure (lifestyles and relative abundances) are important features of Changhsingian brachiopod communities prior to the end-Permian mass extinction (EPME) and could predict temporal and spatial extinction patterns during the EPME. In South China, Changhsingian brachiopod communities show higher diversity than other contemporaneous brachiopod communities in the world and have been reported from a variety of sedimentary environments. In this paper, brachiopods from 18 sections in South China were selected to divide communities and compare their ecological structure. Based on the results of network analysis, cluster analysis and quantitative data from the selected sections, we show that Changhsingian brachiopod communities in South China can be categorized into three assemblages along a marine depth gradient: the Neochonetes–Fusichonetes–Paryphella Assemblage from the shallow-water clastic-rock facies, Spinomarginifera–Peltichia–Oldhamina Assemblage from the shallow-water carbonate platform facies and Fusichonetes–Crurithyris Assemblage from the deep-water siliciclastic intracontinental basin facies. Compared with communities from carbonate platform facies, the communities from siliciclastic facies were characterized by high dominance, low evenness and low lifestyle diversity, which might be important biotic factors leading to earlier extinctions. After the extinction began in all environments, the whole earliest Triassic brachiopod community was first dominated by Fusichonetes and then by Crurithyris. These patterns of domination and replacement could be explained by morphological and ecological advantages. The domination of these two genera, which were already adapted to the oxygen and food-limited deep-water habitat, indicates that the cooler deep-water environment might have been a relatively less stressed habitat after the beginning of the EPME. This suggests that global warming might be the main trigger among the previously proposed synergistic environmental stresses, while anoxia might not, at least for the beginning of EPME.     

我国南沙群岛的植物与植被概况

南沙群岛是我国最南端的一个植物地理区域,但目前我国植物学家对这里的植物与植被还了解得不多。本文通过实地考察个别人工岛,并广泛收集国内外的有关文献资料,概述了南沙群岛的植物与植被的组成、特点及其与邻近地区植物区系的关系,为我国南海岛屿植物区系的研究提供基本资料．     

Neoselachian (Chondrichthyes, Elasmobranchii) diversity across the Cretaceous-Tertiary boundary

Fishes are often thought to have passed through mass extinctions, including the Cretaceous-Tertiary (KT) event, relatively unscathed. We show that neoselachian sharks suffered a major extinction at the K/T boundary. Out of 41 families, 7 became extinct (17±12%). The proportional measure increases at lower taxic levels: 56±10% loss of genera (loss of 60 out of 107) and 84±5% loss of species (loss of 182 out of 216). However, the Maastrichtian and Danian are characterized by a high number of singleton taxa. Excluding singletons we have calculated a 34±11% loss of genera and a 45±9% loss of species. The simple completeness metric (SCM) for genera displays a decrease from the Maastrichtian (94%) to the Danian (85%) indicating a rather complete fossil record of neoselachian genera. The extinctions were heavy among both sharks and batoids (skates and rays), but most severe among batoids, which lost almost all identifiable species. There were equal losses among open marine apex predators (loss of Anacoracidae, Cretoxyrhinidae, and Scapanorhynchidae) and durophagous demersal forms from the continental shelf and shallow seas (Hypsobatidae, Parapaleobatidae, Sclerorhynchidae, Rhombodontidae). Benthopelagic and deep-sea forms were apparently little affected. New families with similar ecological roles (Carcharhinidae, Isuridae, Torpedinidae) replaced these families in the Danian, and full diversity of the different shark and batoid groups had been recovered by the end of the Paleocene or early Eocene. Sharks and rays suffered levels of extinction entirely in line with other groups of organisms at the K/T extinction event.