Paleoecology of Auloporida: an example from the Devonian of the Holy Cross Mts., Poland

Substrate specificity of Auloporida (Tabulata) from the Ska?y Fm. (Upper Eifelian-Lower Givetian) of the Holy Cross Mts., Poland, has been recognized. Kyrtatrypa sp., a rare species in the formation (under 5%), was the most often encrusted brachiopod (59% of investigated specimens), while the most often occurring brachiopod, Aulacella eifeliensis (de Verneuil) was nearly not encrusted. The majority of encrusted brachiopods were larger than 20 mm, while smaller brachiopods occur abundantly in the Formation. The substrate specificity has been caused mainly by the ornamentation of the host's shell. The position of corallites along the commissure of the brachiopod shell proves that auloporids often encrusted living hosts. The epizoan probably used water currents produced by brachiopod's lophophore impoverishing the host's food composition, their relationship can therefore be described as scramble competition.     

Traces of predation/parasitism recorded in Eocene brachiopods from the Castle Hayne Limestone,North Carolina,USA

Schimmel, M., Kowalewski, M. & Coffey, BP. 2011: Traces of predation/parasitism recorded in Eocene brachiopods from the Castle Hayne Limestone, North Carolina, USA. Lethaia, Vol. 45, pp. 274–289. The Castle Hayne Limestone (Middle Eocene, North Carolina), noted for its diverse macro‐invertebrate fossils, was sampled to assess if Early Cenozoic brachiopods from eastern North America record any traces of biotic interactions. Systematic surveys of two North Carolina quarries yielded 494 brachiopods dominated by one species: Plicatoria wilmingtonensis (Lyell and Sowerby, 1845). Despite subtle variations in taphonomy, taxonomy and drilling patterns, the two sampled quarries are remarkably similar in terms of quantitative and qualitative palaeoecological and taphonomic patterns. In both quarries, brachiopods contain frequent drillholes (24.5% specimens drilled). The majority of drillholes were singular, perpendicular to shell surface and drilled from the outside. Ventral valves were drilled slightly more frequently than dorsal ones, but site‐selectivity in drilhole location was not evident. Larger brachiopods were drilled significantly more frequently than smaller ones. However, drillhole diameter did not correlate with brachiopod size. The drillholes are interpreted as records of ‘live‐live’ biotic interactions, representing either predatory attacks or parasitic infestations or a combination of those two types of interactions. A notable fraction of specimens bear multiple drillholes, which is consistent with either parasitic nature of interactions or frequent failed predatory events. The high drilling frequency reported here reinforces other reports (from other continents and other epochs of the Cenozoic), which suggest that brachiopods may be an important prey or host of drilling organisms in some settings. The number of case studies reporting high frequencies of drilling in brachiopods is still limited and thus insufficient to draw reliable generalizations regarding the causes and consequences of these occasionally intense ecological interactions. □Brachiopods, drilling parasitism, drilling predation, Eocene, North Carolina, taphonomy.     

Molecular phylogeny of brachiopods and phoronids based on nuclear-encoded small subunit ribosomal RNA gene sequences

Brachiopod and phoronid phylogeny is inferred from SSU rDNA sequences of 28 articulate and nine inarticulate brachiopods, three phoronids, two ectoprocts and various outgroups, using gene trees reconstructed by weighted parsimony, distance and maximum likelihood methods. Of these sequences, 33 from brachiopods, two from phoronids and one each from an ectoproct and a priapulan are newly determined. The brachiopod sequences belong to 31 different genera and thus survey about 10% of extant genus-level diversity. Sequences determined in different laboratories and those from closely related taxa agree well, but evidence is presented suggesting that one published phoronid sequence (GenBank accession UO12648) is a brachiopod-phoronid chimaera, and this sequence is excluded from the analyses. The chiton, Acanthopleura, is identified as the phenetically proximal outgroup; other selected outgroups were chosen to allow comparison with recent, non-molecular analyses of brachiopod phylogeny. The different outgroups and methods of phylogenetic reconstruction lead to similar results, with differences mainly in the resolution of weakly supported ancient and recent nodes, including the divergence of inarticulate brachiopod sub-phyla, the position of the rhynchonellids in relation to long- and short-looped articulate brachiopod clades and the relationships of some articulate brachiopod genera and species. Attention is drawn to the problem presented by nodes that are strongly supported by non-molecular evidence but receive only low bootstrap resampling support. Overall, the gene trees agree with morphology-based brachiopod taxonomy, but novel relationships are tentatively suggested for thecideidine and megathyrid brachiopods. Articulate brachiopods are found to be monophyletic in all reconstructions, but monophyly of inarticulate brachiopods and the possible inclusion of phoronids in the inarticulate brachiopod clade are less strongly established. Phoronids are clearly excluded from a sister-group relationship with articulate brachiopods, this proposed relationship being due to the rejected, chimaeric sequence (GenBank UO12648). Lineage relative rate tests show no heterogeneity of evolutionary rate among articulate brachiopod sequences, but indicate that inarticulate brachiopod plus phoronid sequences evolve somewhat more slowly. Both brachiopods and phoronids evolve slowly by comparison with other invertebrates. A number of palaeontologically dated times of earliest appearance are used to make upper and lower estimates of the global rate of brachiopod SSU rDNA evolution, and these estimates are used to infer the likely divergence times of other nodes in the gene tree. There is reasonable agreement between most inferred molecular and palaeontological ages. The estimated rates of SSU rDNA sequence evolution suggest that the last common ancestor of brachiopods, chitons and other protostome invertebrates (Lophotrochozoa and Ecdysozoa) lived deep in Precambrian time. Results of this first DNA-based, taxonomically representative analysis of brachiopod phylogeny are in broad agreement with current morphology-based classification and systematics and are largely consistent with the hypothesis that brachiopod shell ontogeny and morphology are a good guide to phylogeny.     

STROPHOMENIDE BRACHIOPODS FROM THE CHANGWU FORMATION (LATE KATIAN, LATE ORDOVICIAN) OF CHUN'AN, WESTERN ZHEJIANG, SOUTH-EAST CHINA

Abstract:  Late Ordovician strophomenide brachiopods (superfamilies Strophomenoidea and Plectambonitoidea) from the upper Changwu Formation (mid Ashgill, late Katian) of Jianglütang, Chun'an County, western Zhejiang Province, consist of ten genera and 12 species. Five new species of three new genera are recognized: Chunanomena triporcata , Chunanomena sembellina , Cheramomena subsolana , Lateriseptomena modesta , and Lateriseptomena rugosa . The strophomenide brachiopods from the upper Katian strata described in this study and those from the border region of Zhejiang and Jiangxi provinces reported in previous work contain 16 strophomenoid and 12 plectambonitoid genera, and most of the strophomenoids are endemic to South China. Numerical analysis of well-documented late Katian strophomenide brachiopod faunas indicates a strong provincialism, characterized by the highly distinct North American province (Laurentia), the South China-Kazakhstan province, and the Avalonia-Baltica province (Wales, Belgium and Sweden). Surprisingly, the Girvan district of Scotland, which was a peri-Laurentian terrane during the Ordovician, contains a late Katian brachiopod fauna that is more closely related to the contemporaneous brachiopods of Avalonia-Baltica than to those of North America.     

WUCHIAPINGIAN (LOPINGIAN, LATE PERMIAN) BRACHIOPODS FROM THE EPISKOPI FORMATION OF HYDRA ISLAND, GREECE

Abstract:  Twenty-three species of silicified brachiopods are described from four samples in the middle and upper parts of the Episkopi Formation from Hydra Island, Greece. These brachiopods are newly recorded from the region and together with previously described brachiopods from the same localities constitute the most diverse Lopingian (Late Permian) brachiopod fauna reported in southern Europe. The brachiopod fauna is Wuchiapingian as indicated by the associated conodonts. The fauna from Hydra exhibits strong palaeobiogeographical links with the faunas from South China. In addition, palaeobiogeographical affinities with the faunas of Thailand and the northern peri-Gondwanan region are also present, which implies a peri-Gondwanan origin for Hydra. Palaeoecologically, the brachiopod assemblage from sample EP in the middle part of the Episkopi Formation is dominated by pedically-attached and cementing genera and reflects moderate energy conditions above storm wave base and an abundance of hard substrates provided by sponges in the biohermal habitat. By contrast, the brachiopod assemblage in the other three samples from the upper part of the Episkopi Formation is dominated by spinose genera with a free-resting life habit, suggesting soft substrates in a quiet water environment below storm wave base on the outer part of the shelf. New taxa are Petinospiriferina gen. nov., Hustedia episkopiensis sp. nov., Waterhouseiella hydraensis sp. nov. and Xenosaria tenuis sp. nov.     

PHYLOGENETIC RELATIONSHIPS AMONG EXTANT BRACHIOPODS

Abstract— The monophyletic status of the Brachiopoda and phylogenetic relationships within the phylum have long been contentious issues for brachiopod systematists. The relationship of brachiopods to other lophophore-bearing taxa is also uncertain; results from recent morphological and molecular studies are in conflict. To test current hypotheses of relationship, a phylogenetic analysis was completed (using PAUP 3.1.1) with 112 morphological and embryological characters that vary among extant representatives of seven brachiopod superfamilies, using bryozoans, phoronids, pterobranchs and sipunculids as outgroups. In the range of analyses performed, brachiopod monophyly is well supported, particularly by characters of soft anatomy. Arguments concerning single or multiple origins of a bivalved shell are not relevant to recognizing brachiopods as a clade. Articulate monophyly is very strongly supported, but inarticulate monophyly receives relatively weak support. Unlike previous studies, the nature of uncertainties about the clade status of Inarticulata are detailed explicitly here, making them easier to test in the future. Calcareous inarticulates appear to share derived characters with the other inarticulates, while sharing many primitive characters with other calcareous brachiopods (the articulates). Experimental manipulation of the data matrix reveals potential sources of bias in previous hypotheses of brachiopod phylogeny. Although not tested explicitly, lophophorate monophyly is very tentatively supported. Molecular systematic studies of a diverse group of brachiopods and other lophophorates will be particularly welcome in providing a test of the conclusions presented here.     

Molecular paleobiological insights into the origin of the Brachiopoda

Most studies of brachiopod evolution have been based on their extensive fossil record, but molecular techniques, due to their independence from the rock record, can offer new insights into the evolution of a clade. Previous molecular phylogenetic hypotheses of brachiopod interrelationships place phoronids within the brachiopods as the sister group to the inarticulates, whereas morphological considerations suggest that Brachiopoda is a monophyletic group. Here, these hypotheses were tested with a molecular phylogenetic analysis of seven nuclear housekeeping genes combined with three ribosomal genes. The combined analysis finds brachiopods to be monophyletic, but with relatively weak support, and the craniid as the sister taxon of all other brachiopods. Phylogenetic-signal dissection suggests that the weak support is caused by the instability of the craniid, which is attracted to the phoronids. Analysis of slowly evolving sites results in a robustly supported monophyletic Brachiopoda and Inarticulata (Linguliformea+Craniiformea), which is regarded as the most likely topology for brachiopod interrelationships. The monophyly of Brachiopoda was further tested with microRNA-based phylogenetics, which are small, noncoding RNA genes whose presence and absence can be used to infer phylogenetic relationships. Two novel microRNAs were characterized supporting the monophyly of brachiopods. Congruence of the traditional molecular phylogenetic analysis, microRNAs, and morphological cladograms suggest that Brachiopoda is monophyletic with Phoronida as its likely sister group. Molecular clock analysis suggests that extant phoronids have a Paleozoic divergence despite their conservative morphology, and that the early brachiopod fossil record is robust, and is not affected by taphonomic factors relating to the late-Precambrian/early-Cambrian phosphogenic event.     

Brachiopods, biostratigraphy, and correlation of the Permian marine deposits of Mongolia

Six horizons and 11 brachiopod zones are distinguished in the Permian of the Khangai-Khentei (Boreal) and South Mongolian (Tethys) marine basins of Mongolia on the base of monographic study of brachiopods and literary data on other faunal groups. The distinguished in the Permian basins of Mongolia biostratons are correlated with zonal brachiopod scales of neighboring regions of Northeast Asia. Possible bio-geographical relations of the Mongolian Permian marine basins and channels of brachiopods are analyzed. Forty-five brachiopod species are figured including species, which were first described from Mongolia, zonal index species and typical representatives of zonal assemblage.     

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。     

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.     

Recent brachiopods from the southern Brazilian shelf: palaeontological and biogeographical implications

Until recently, the rhynchonelliform (articulated) brachiopod fauna from the Brazilian continental shelf (western South Atlantic) was represented only by the endemic species Bouchardia rosea (Mawe), reported from coastal waters of the states of São Paulo and Rio de Janeiro. The present study, based on samples from coastal (<30 m), shelf, and continental slope waters (99–485 m), documents the South Atlantic brachiopod fauna and shows that this fauna is more widespread, diverse, and cosmopolitan than previously thought. Based on a total of 16,177 specimens, the following brachiopods have been identified: Bouchardia rosea (Family Bouchardiidae), Platidia anomioides (Family Platidiidae), Argyrotheca cf. cuneata (Family Megathyrididae), and Terebratulina sp. (Family Cancellothyrididae). In coastal settings, the fauna is overwhelmingly dominated by Bouchardia rosea . Rare juvenile (<2 mm) specimens of Argyrotheca cf. cuneata were also found at two shallow-water sites. In shelf settings (100–200 m), the fauna is more diverse and includes Bouchardia rosea , Terebratulina sp., Argyrotheca cf. cuneata , and Platidia anomioides . Notably, Bouchardia rosea was found in waters as deep as 485 m, extending the known bathymetric range of this genus. Also, the record of this brachiopod in waters of the state of Paraná is the southernmost known occurrence of this species. The genera Platidia and Terebratulina are documented here for the first time for the western South Atlantic. The Brazilian brachiopod fauna shares similarities with those from the Atlantic and Indian shelves of southern Africa, and from the Antarctic, Caribbean and Mediterranean waters. The present-day brachiopods of the western South Atlantic are much more cosmopolitan than previously thought and their Cenozoic palaeobiogeographic history has to be reconsidered from that perspective.     

Ecological interactions on macroevolutionary time scales: clams and brachiopods are more than ships that pass in the night

Competition among organisms has ecological and evolutionary consequences. However, whether the consequences of competition are manifested and measureable on macroevolutionary time scales is equivocal. Marine bivalves and brachiopods have overlapping niches such that competition for food and space may occur. Moreover, there is a long‐standing debate over whether bivalves outcompeted brachiopods evolutionarily, because brachiopod diversity declined through time while bivalve diversity increased. To answer this question, we estimate the origination and extinction dynamics of fossil marine bivalve and brachiopod genera from the Ordovician through to the Recent while simultaneously accounting for incomplete sampling. Then, using stochastic differential equations, we assess statistical relationships among diversification and sampling dynamics of brachiopods and bivalves and five paleoenvironmental proxies. None of these potential environmental drivers had any detectable influence on brachiopod or bivalve diversification. In contrast, elevated bivalve extinction rates causally increased brachiopod origination rates, suggesting that bivalves have suppressed brachiopod evolution.     

Pennsylvanian sea level cycles, nutrient availability and brachiopod paleoecology

As with modern organisms, the spatial and temporal distribution of fossil communities was controlled by both the physical setting in which the organisms lived and by the organisms' physiology and interactions. By studying the sedimentological and geochemical context of fossil communities, it is possible to assess the relative importance of the physical setting and the organisms' physiology. Comparison of Pennsylvanian brachiopod associations with changing sedimentological context (water depth/facies) and nutrient availability indicates that body size is a function of water depth and nutrient availability for most spire-bearing (athyridids and spiriferids) brachiopods but rarely for productid brachiopods. Spire-bearing brachiopods dominate the associations in high-nutrient settings, and productid brachiopods dominate the associations in low-nutrient settings. This difference suggests that physiological differences between brachiopod orders, such as lophophore filtering efficiency, play an important role in controlling their distribution.     

The life and death of torquirhynchia inconstans (brachiopoda,upper jurassic) in England

Torquirhynchia inconstans, a rhynchonellid brachiopod, shows a curious asymmetric commissure. This is interpreted as an adaptation to life in tidal environments. Type of preservation, growth line, and size frequency analysis indicate that the population analyzed consist of dominantly mature individuals, which by analogy with recent brachiopod populations is a primary feature of the original population, and not due to selective destruction, selective transport, or selective predation of smaller individuals. The possible functions of the asymmetry of Torquirhynchia inconstans are considered, and it is concluded that the brachiopod was adapted to life in tidal environments, a conclusion supported by sedimentological evidence. Asymmetric brachiopods are considered to have developed from colonies of partly asymmetric, variable brachiopods by selection of extreme variants.     

Functional disposition of the lophophore in living Brachiopoda

Emig, C. C. 1992 07 15: Functional disposition of the lophophore in living Brachiopoda.
The shape and disposition of adult brachiopod lophophores relate to in- and excurrent apertures. to the internal water irrigation system, to shell orientation at substratum and to near-bottom currents. The arrangement of the mantle canals and gonads of different lophophores are induccd by water circulation. The trocholophe (2% of living species) is considered as a plesiomorphic character which represents the basic plan of the lophophore, shared by all Lophophorata. Three different types of schizolophe (10%) are represented in terebratuloids, thecidioids and discinids. The spirolophe (19%), characteristic of rhynchonellides and most inarticulate brachiopods, except the schizolophe Pelagodiscus , has evolved divergently into specific arrangements of the mantle canals and gonads. The zygo-plectolophe (67%) is characteristic of most Terebratulida. The ptycholophe (2%) probably evolved independently in Megathlris and the thecidioids. The mesolophe, known in the fossil chonetdceans, is considered to be a primitive zygo-plectolophe. The median brachiopod sulcus increases the efficiency of the excurrent system and is considered as an evolved character but a homoplasy within the brachiopods. The characteristics of Recent lophophore types have to be taken into account when reconstructing the lophophore in fossil forms. Brachiopoda, Lophophorata, lophophore, water system, orientation, evolution .     

A brachiopod boring (Podichnus) in a Late Cretaceous oyster from a mangrove-like environment, Skåne, Sweden

A characteristic etching trace, comparable to those produced by pedicles of some Recent brachiopods, was found on a Late Cretaceous, Campanian oyster shell from a mangrove-like setting in the Åsen quarry, Sweden. The trace fossil belongs to the ichnospecies Podichnus centrifugalis Bromley & Surlyk, 1973 and is, together with a single specimen of Crania craniolaris Linnæus, 1758, the only evidence of brachiopods in the mangrove-like environment.     

A new morphometric model in distinguishing two closely related extinct brachiopod species

Pseudoatrypa (Atrypida, Brachiopoda) are extinct Paleozoic brachiopods that are the closest relatives of the modern brachiopods, rhynchonellids. Two Paleozoic brachiopod species were qualitatively distinguished based on close examination of phenotypic characters and traditional morphometrics. These species were recovered for the first time from the Givetian localities of Traverse Group of Michigan. The efficacy of geometric morphometric analyses in discriminating species has been proven for various organisms, but rarely studies have been performed in distinguishing brachiopod species. The advanced morphometric model developed here using the Procrustes-based landmark method is based on the analysis of valve shape in two extinct brachiopod species: Pseudoatrypa lineata and Pseudoatrypa devoniana. Results from quantitative analyses reinforce the qualitative identifications of the two species. The application of this method on four different orientations (dorsal, ventral, posterior and anterior regions) of the brachiopods permits the two species to be clearly separated, and this study proposes that the model can be effectively used for future identification and distinction of fossil species. Additionally, this study will aid biologists, paleontologists and neontologists in making appropriate advancement in systematic studies and further assist with gauging biodiversity congruously.     

Evolutionary significance of a middle Cambrian (Series 3) in situ occurrence of the pedunculate rhynchonelliform brachiopod Nisusia sulcata

Exceptionally preserved, silicified and articulated complete shells of the rhynchonelliform kutorginate brachiopod Nisusia sulcata are redescribed from the middle Cambrian (Series 3) Marjum Limestone, Utah. Cylindroid sausage‐like protrusions, emerging posteriorly between the valves, were originally interpreted as faecal in origin, but restudy under the SEM shows that these features represent silicified pedicles as they are attached in situ to other Nisusia. The Nisusia host most likely was alive at the time of attachment. Restudy of the pedicles of Nisusia provides new phylogenetic information on the anatomy of the earliest rhynchonelliforms. The silicified pedicles differ considerably from the pedicles of living crown group rhynchonelliforms in being strongly annulated, distally tapering, and were likely to have been rather more flexible. The Nisusia pedicles are more similar to the exceptionally preserved pedicles from other Cambrian rhynchonelliform brachiopods, including Kutorgina, Longtancunella and Alisina, but these emerge from the ventral apical foramen rather than from between the valves as in Nisusia. Although generally similar, these two types of pedicles are unlikely to represent homologous structures as Nisusia is provided with both an apical foramen (possibly larval attachment) and a posterior adult pedicle. The similarities may be explained by similar type of accretionary growth from two different types of epithelia. The Nisusia‐like pedicle appeared early within the kutorginates and rhynchonellates. The discovery of hollow spines in Nisusia sulcata further supports the generic assignation of the species.     

Supraordinal brachiopod classification

Three controversial problems of brachiopod supraordinal classification are discussed: the position of brachiopods in the classification of Metazoa, their classification at phylum and class level, and the classification of the articulate brachiopods. The position of brachiopods in the system of Metazoa remains uncertain. There are no strong reasons for changing the traditional division of the phylum Brachiopoda into the classes Inarticulata and Articulata. The class Articulata is divided into the subclasses Orthata, Strophomenata, Spiriferata, and Terebratulata.     

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.