Diversity and extinction patterns of Permian coral faunas of China

Coral diversity and extinction patterns in the Permian of China are revealed through statistical analyses of 56 coral families, 263 genera and 2100 species from five consecutive time intervals. The highest coral diversity is in the Chuanshanian, with 753 species, 167 genera and 39 families. In contrast, the lowest diversity is in the Changhsingian, with only 68 species, 20 genera and 10 families. Two decreases in diversity can be recognized during the Permian. The first occurred at the end of the Maokouan (end-Guadalupian) and is marked by the loss of 75.6% of coral families, 77.8% of coral genera and 82.2% of coral species. The second major diversity drop took place at the end of the Changhsingian, when all rugose and tabulate corals became extinct. The extinction at the end of the Guadalupian in Pangea may be related to the middle Permian global regression. However, in South China the end-Maokouan extinction may be related to the eruption of the Omeishan Basalt. A triple-zoned palaeobiogeographical pattern is well expressed by coral diversity.     

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.     

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).     

茅口期相对海平面变化对类动物群的影响——以贵州盘县火铺镇茅口组剖面为例

分析贵州盘县火铺镇茅口期海平面变化的过程和Ｔｉｎｇ类动物群的变化规律,研究相对海平面变化对Ｔｉｎｇ类动物群构及演替的影响。相对海平面的升降变化可引起水深及相关生态条件的改变,并对Ｔｉｎｇ类动物群产生影响,相对海平面变化的幅度和速率是造成Ｔｉｎｇ类动物群面貌变化的重要因素,三级相对海平面上升末期和下降未期,海平面的大幅度抬升或下降是造成Ｔｉｎｇ类物种丰度下降,新种增加量减少的重要原因,海平面的持续缓     

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.     

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。     

Latest Guadalupian brachiopods from the Guadalupian/Lopingian boundary GSSP section at Penglaitan in Laibin, Guangxi, South China and implications for the timing of the pre-Lopingian crisis

A brachiopod fauna comprising nine species in eight genera from three closely spaced stratigraphic horizons of the same stratigraphic section is described for the first time from the Laibin Limestone in the uppermost part of the Maokou Formation in the Guadalupian/Lopingian (G/L) GSSP section at Penglaitan, Guangxi Autonomous Region, South China. The brachiopod assemblages are bracketed between two conodont zones: Jinogondolella xuanhanensis Zone below and Jinogondolella granti Zone above and, therefore, they can be safely assigned to the latest Capitanian in age. However, all but one of the nine brachiopod species from the Laibin Limestone carry strong early Lopingian (Wuchiapingian) aspect. Thus, the discovery of this brachiopod fauna not only suggests that some Lopingian brachiopod species had already appeared in the late Guadalupian (Capitanian); more importantly, it has also highlighted the fact that both the previously noted pre-Lopingian life crisis (or end-Guadalupian or Middle Permian mass extinction) and Lopingian recovery/radiation actually occurred in late Capitanian times, sometime before the G/L chronostratigraphic boundary. So far, the Penglaitan GSSP section provides the highest-resolution disappearance patterns of different fossil groups around the G/L boundary.     

Cancrinella and Costatumulus (Brachiopoda) from the Permian of South Mongolia and South China: Their morphology,biostratigraphy and distribution

Two brachiopod genera, Cancrinella Fredericks and Costatumulus Waterhouse, are studied from the Permian of South Mongolia and South China. Many of the specimens previously described as Cancrinella are revised and assigned to Costatumulus. The new data presented in this paper indicate that these two genera are similar to each other, but are distinguishable by different features in relation to their morphology. Several species of both genera are described here, providing critical new information on the morphology and taxonomy of Cancrinella and Costatumulus. The reported biostratigraphical information reveals that species of Cancrinella are present from the Lower Carboniferous (Tournaisian) to Middle Permian (Capitanian), whereas species of Costatumulus are mostly restricted to the Lower Permian (Sakmarian) to Upper Permian (Changhsingian). An analysis of the palaeogeographical distribution of Cancrinella and Costatumulus reveals that Cancrinella was more commonly distributed in the Northern Transitional Zone and the Boreal Realm, in contrast to Costatumulus, which tended to be more common in the Southern Transitional Zone and Gondwanan Realm.     

Late Permian marine ecosystem collapse began in deeper waters: evidence from brachiopod diversity and body size changes

Analysis of Permian–Triassic brachiopod diversity and body size changes from different water depths spanning the continental shelf to basinal facies in South China provides insights into the process of environmental deterioration. Comparison of the temporal changes of brachiopod diversity between deepwater and shallow‐water facies demonstrates that deepwater brachiopods disappeared earlier than shallow‐water brachiopods. This indicates that high environmental stress commenced first in deepwater settings and later extended to shallow waters. This environmental stress is attributed to major volcanic eruptions, which first led to formation of a stratified ocean and a chemocline in the outer shelf and deeper water environments, causing the disappearance of deep marine benthos including brachiopods. The chemocline then rapidly migrated upward and extended to shallow waters, causing widespread mass extinction of shallow marine benthos. We predict that the spatial and temporal patterns of earlier onset of disappearance/extinction and ecological crisis in deeper water ecosystems will be recorded during other episodes of rapid global warming.     

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.     

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。     

Callistophytalean pteridosperms from Permian aged floras of China

Abstract: Recent investigations into Permian aged floras from China have highlighted the widespread occurrence of callistophytalean pteridosperms that challenge previous understanding of their spatial and temporal distribution and diversity. In China, the group spans the Permian period and constitutes a distinctive but rare component in many peat‐forming environments. The stratigraphically earliest callistophytalean occurs in the Asselian‐Sakmarian stages with fossils from the Taiyuan Formation of northern China including ovules of Callospermarion undulatum in coal ball assemblages, and ovulate fronds of Norinosperma shanxiensis and synangiate fronds of Norinotheca shanxiensis in adpression assemblages. More abundant in the fossil records are adpression remains from the Roadian‐Wordian stages with the Lower Shihhotse Formation preserving abundant vegetative and ovulate remains of Emplectopteris triangularis that is now considered to represent a callistophytalean. The youngest callistophytalean recognised is from the Wuchaipingian‐Changhsingian stages with the Xuanwei Formation of southern China containing a single stem of Callistophyton boyssetii that provides indisputable evidence of the group in the lead up to the end‐Permian mass extinction. These accounts are augmented by analysis of pollen records that demonstrate the callistophytalean pollen genus Vesicaspora to be widespread through palynological assemblages from the Permian period in both North and South China, including the Upper Shihhotse Formation, Shihchienfeng Group, Xuanwei Formation, and possibly also in the mid‐Pennsylvanian Benxi Formation. Although macrofossil specimens are uncommon elements in the assemblages that contain them, they demonstrate the continuity of callistophytalean pteridosperms from the Pennsylvanian sub‐period into the early Guadalupian epoch of the Permian in North China and into the Lopingian epoch of the Permian in South China. Of the species present, both Callistophyton boyssetii and Callospermarion undulatum are known from the Pennsylvanian–earliest Permian age floras of Euramerica, whereas Norinosperma, Norinotheca and Emplectopteris appear to represent endemic Cathaysian elements. Results imply that callistophytalean pteridosperms can no longer be excluded from theories of post‐Carboniferous plant evolution and floristics, appearing to have played an important role in both Permian and Carboniferous aged plant communities. The presence of Vesicaspora in several formations from which macro‐remains have not been identified is a hopeful indicator that further callistophytalean pteridosperms are yet to be found.     

Silurian craniide brachiopods from Gotland

Seven craniide brachiopod genera are described from the Silurian (Wenlock–Ludlow) of Gotland, including one new genus and five new species. The new genus and species Thulecrania septicostata is unique among Silurian craniides as it possesses solid spines. The new species Lepidocrania multilamellosa is the first Silurian record of this poorly known Permian genus. The problematic North American Propatella Grubbs, 1939 , was originally described as a gastropod, but the new species Propatella palmaria from Gotland shows that it is a craniide with sutured hollow spines of a type not previously recorded from Silurian craniides. The dorsal valves of the new species Valdiviathyris? bicornis are remarkably similar to those of the type species and represent the first possible Palaeozoic record of this poorly known extant craniide. This first systematic study of craniide brachiopods from the Silurian of Gotland shows that the diversity is relatively high as compared to other known Silurian craniide faunas, but a more thorough comparison is not possible due to the lack of data from most parts of the world. The new data from Gotland support the view that the craniides were not affected by the end‐Ordovician extinction.     

Permophricodothyris Pavlova, 1965 (Brachiopoda, Spiriferida) from the Permian of South China: its morphology, biostratigraphy and distribution

A study of both silicified and nonsilicified specimens of Permian reticularioid brachiopods from South China suggests thatPermophricodothyris, a genus previously rarely reported from China, is actually very common and abundant in the Middle and especially Upper Permian of South China. This study also clarifies, for the first time, that many of the reticularioid brachiopod species previously described asSquamularia in fact belong toPermophricodothyris. The new data presented in this paper also allows a critical evaluation ofPermophricodothyris in relation to its closest allies:Phricodothyris, Squamularia, Bullarina andNeophricodothyris. The revision reveals that a total of 18Permophricodothyris species are present in the Middle and Upper Permian of South China, with only one species,P. squamularioides, having survived the Permian-Triassic mass extinction. Two species,P. grandis (Chao) andP. guangxiensis Han, Zhou & Wang, are redescribed here, providing critical new information on the morphology and taxonomy of these species.     

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.     

Large extinctions of articulate brachiopods in the paleozoic and their ecological and evolutionary consequences

The largest Paleozoic extinctions of articulate brachiopods occurred at the Frasnian—Famennian boundary in the Late Devonian and at the Permian—Triassic boundary. Both extinctions affected taxa of all levels, including orders, but differed in scale, course, and ecological and evolutionary consequences. The Frasnian—Famennian extinction event was selective and evolutionary activity after the crisis varied in different orders. However, in the Early Carboniferous, the brachiopod diversity was mostly restored in comparison with the Devonian maximum. In particular groups, preadaptation played a role in changes in diversity and reconstruction of communities. The brachiopod composition at this boundary changed sharply. The extinction event at the end of Permian was global and accompanied by changes in the biota. Later, in the Meso-Cenozoic, the brachiopod diversity was not restored, and bivalves acquired primary importance in various bottom communities of different sea zones where Paleozoic brachiopods previously dominated. Extinction of brachiopods at this boundary was long and gradual. The symptoms of the ecological crisis in the development of Permian brachiopods are recognized beginning from the Roadian Age, which was probably the onset of this crisis.     

Latitudinal diversity gradients for brachiopod genera during late Palaeozoic time: links between climate, biogeography and evolutionary rates

Aim The latitudinal diversity gradient, in which taxonomic richness is greatest at low latitudes and declines towards the poles, is a pervasive feature of the biota through geological time. This study utilizes fossil data to examine how the latitudinal diversity gradient and associated spatial patterns covaried through the major climate shifts at the onset and end of the late Palaeozoic ice age. Location Data were acquired from fossil localities from around the world. Methods Latitudinal patterns of diversity, mean geographical range size and macroevolutionary rates were constructed from a literature‐derived data base of occurrences of fossil brachiopod genera in space and time. The literature search resulted in a total of 18,596 occurrences for 991 genera from 2320 localities. Results Climate changes associated with the onset of the late Palaeozoic ice age (c. 327 Ma) altered the biogeographical structure of the brachiopod fauna by the preferential elimination of narrowly distributed, largely tropical genera when glaciation began. Because the oceans were left populated primarily with widespread genera, the slope of the diversity gradient became gentle at this time, and the gradient of average latitudinal range size weakened. In addition, because narrowly distributed genera had intrinsically high rates of origination and extinction, the gradients of both of these macroevolutionary rates were also reduced. These patterns were reversed when the ice age climate abated in early Permian time (c. 290 Ma): narrowly distributed genera rediversified at low latitudes, restoring steep gradients of diversity, average latitudinal range size and macroevolutionary rates. Main conclusions During late Palaeozoic time, these latitudinal gradients for brachiopods may have been linked by the increased magnitude of seasonality during the late Palaeozoic ice age. Pronounced seasonality would have prevented the existence of genera with narrow latitudinal ranges. These results for the late Palaeozoic ice age suggest a climatic basis for the present‐day latitudinal diversity gradient.     

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。     

Conodonts from the Early Triassic microbialite of Guangxi (South China): implications for the definition of the base of the Triassic System

We describe a new Early Triassic (Griesbachian) succession of conodont faunas from a high‐resolution sampling of the basal Early Triassic microbial limestone and the base of the overlying unit at the Wuzhuan section (Nanpanjiang Basin, Guangxi, South China). The microbial limestone records the earliest phase of the Early Triassic biotic recovery after the end‐Permian mass extinction. For the first time, rich conodont faunas are reported from within the microbialite. The faunas from Wuzhuan are largely dominated by anchignathodontids, including several Isarcicella species, which were previously documented only from strata above the microbialite. A total of 14 conodont species assigned to three genera is recorded from the Wuzhuan section. Starting from the base of the microbialite upwards, several species are sequentially added to the conodont assemblage. The alpha diversity peaks at the top of the microbialite. The conodont record in the considered microbialite interval at Wuzhuan is presumably unaffected by local ecological changes. It therefore more likely represents an evolutionary rather than an ecological pattern. We compare the Wuzhuan's conodont record with a well‐supported phylogenetic model and suggest that the sequence of first occurrences at Wuzhuan is the closest to the ‘true’ sequence of evolutionary events that took place during this Griesbachian radiation of anchignathodontids. Based on comparisons with the GSSP section at Meishan, we suggest further that the first occurrence of Hindeodus parvus in Meishan does not correspond to its first appearance datum.     

Global diversity and endemism in Early Silurian (Aeronian) brachiopods

Silurian brachiopods of Middle Llandovery (Aeronian) ages are reviewed, and 215 genera are identified here, compared with 109 in the preceding Early Llandovery (Rhuddanian), indicating a recovery‐radiation interval after the end‐Ordovician mass extinction. The chief regions in which they are found are the continents of South China, Avalonia‐Baltica, Laurentia and Siberia, which were all at tropical latitudes with the exception of Avalonia. In addition, the very large superterrane of Gondwana, although with patchy brachiopod distribution, included temperate faunas, as well as subtropical faunas (in Iran and Afghanistan). Aeronian brachiopods greatly increased in diversity, with dominance of four major groups: orthides and strophomenides (which had flourished previously in the Rhuddanian), pentamerides and atrypides (which became dominant in the Aeronian), and many newly evolved taxa, and occupied deeper water and wider ecological niches (level bottom and reef) than those in the Rhuddanian. Each of the continents has some endemic genera, but there is a greater proportion of them in South China, where some groups (such as the pentamerides and atrypides) are more diverse and others, such as the orthides, are much less common than elsewhere. Affinity indices (AI) show that two megaunits can be recognized: South China and Avalonia‐Baltica‐Laurentia (ABL); Siberia seems to have been loosely connected with ABL and even more loosely to South China presumably because of its geographical separation in the Northern Hemisphere. The separation of South China from the other megaunits is further supported by cluster analysis.