δO and δC of Permian brachiopods: A record of seawater evolution and continental glaciation

Two hundred sixty-four δ¹⁸O and δ¹³C values of Permian articulate brachiopod shells were analyzed and 81 of these values were characterized as well preserved and biostratigraphically well defined. These were then utilized for construction of baseline oxygen and carbon isotope curves for the Permian interval. In addition, 21 δ¹³C whole rock values are reported for the Wordian and Capitanian.The early Permian, Asselian to Artinskian, times are characterized by ∼ 2.5‰ decrease in oxygen isotope values, from ∼ − 0.7‰ to − 3.3‰ (V-PDB). This is attributed to a ∼ 4-7 °C increase in temperature in the Southern Urals, concomitant with the retreat of the Permo-Carboniferous ice sheets and return of the ¹⁸O-depleted melt water into the oceans. The Late Permian samples from Iran (Jolfa at Kuh-e-Ali-Bashi) and China (Meishan) yield δ¹⁸O values, and presumably temperatures, similar to those that followed the termination of the large-scale glaciation in the Lower Permian. In between, the upper Kungurian to Capitanian samples from the Delaware Basin (Guadalupe Mountains) are enriched in ¹⁸O, at − 1.5‰ to − 3‰. We have no definitive explanation for these data, but tentatively suggest that the “anomaly” can potentially be a result of evaporative enrichment of seawater in ¹⁸O, due to intracratonic arid setting of the basin. The ¹⁸O-enriched nature of the Zechstein samples (− 1.2‰ to + 2.5‰), on the other hand, is in all probability a reflection of the high evaporation rates in the Zechstein sea.The Permian interval is characterized by a relatively constant δ¹³C, at about 4‰. The exceptions are again the brachiopods from the Delaware Basin (Guadalupe Mountains), which show ∼ 1.6‰ increase in the Guadalupian, to values of up to 5.9‰ in the Wordian. A tentative explanation, as in the case of oxygen, is based on the proposition that the semi-enclosed Delaware Basin was likely stratified, with sequestration of the ¹³C-depleted carbon to the deeper water layers and a complementary ¹³C enrichment in the upper oxygenated layer. The coeval open ocean water DIC may have been similar to that of the remainder of the Permian interval, at ∼ 4‰, as indicated by whole rock carbonate samples from Oman, Sicily, and Iran. In the latest Permian, the trend mimics the well-known δ¹³C drop at the Permian/Triassic boundary.     

Surviving the end-Ordovician extinctions: evidence from the earliest Silurian brachiopods of northeastern Jiangxi and western Zhejiang provinces, East China

Earliest Rhuddanian (Silurian) brachiopods are recorded from the basal part of the Lower Llandovery Shiyang and Anji formations in western Zhejiang and northeastern Jiangxi provinces, East China. Associated graptolites including Normalograptus jerini indicate the lowest Rhuddanian Akidograptus ascensus Biozone. The surviving brachiopod fauna includes 19 genera dominated by orthids and strophomenids, whereas pentamerids and atrypids that inhabited mainly warmer water regimes, and were almost absent in the cool/cold Hirnantia Fauna, occur rarely in the studied fauna. Each family is represented by a single genus that seeded their recovery. The predominance of these long-ranging and widely distributed genera is one of major characters of the brachiopod survival in east China. From qualitative and quantitative analysis of faunal composition, diversity and abundance, with evidences from palaeoecology and palaeogeography, the Levenea qianbeiensis Association, Katastrophomena-Leptaena-Levenea Association, and Glyptorthis-Epitomyonia-Levenea Association are recognized and assigned to BA (Benthic Assemblage) 2, BA3, and an ecozone close to the BA3-4 boundary respectively. No Lazarus genera are recorded in this study. Skenidioides and Epitomyonia were chiefly regarded as deeper-water taxa in the Ordovician and Silurian, but are recorded from shallow-water in east China during the early Rhuddanian, indicating an ecologic experiment with these taxa migrating from deep into shallower, better-oxygenated sites at the crisis time and during the subsequent survival interval. This study further demonstrates that the brachiopod faunal turnover after the end-Ordovician extinctions may not have been completed until the late Rhuddanian in South China.     

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.     

广西中泥盆世腕足动物空间分布及古环境意义

统计广西中泥盆世不同沉积相区二十条剖面的腕足动物化石记录共计13目45科69属167种,其中Eifelian期47属73种,Givetian期32属100种.对不同级别分类单元的多样性分析表明,从Eifelian期到Givetian期,不仅腕足动物各个目的组成及多样性结构有明显改变,而且各个时期不同沉积环境下的腕足动物...     

Shell structure, ontogeny and affinities of the Lower Cambrian bivalved problematic fossil Mickwitzia muralensis Walcott, 1913

Exceptionally preserved carbonate- and shale-hosted Mickwitzia muralensis from the Lower Cambrian Mural Formation, southern Canadian Rocky Mountains, complement one another to yield an unusually complete account of its ontogeny, ecology and phylogenetic relationships. The shell of M. muralensis is composed of dense phosphatic layers interspersed with porous organic-rich layers. At the insertion of shell-penetrating tubes, shell layers deflect inwards to produce inwardly pointing cones. The tubes are interpreted as having hosted setae that were secreted by outer-epithelial follicles. Follicular setae also occurred at the mantle margin, where they were oriented within the plane of the shell as in modern brachiopods. During ontogeny, the initial setae oriented in the plane of the shell occurred before the first shell-penetrative setae. In the juvenile and early-mature stages of shell secretion, a posterior opening was present between both valves and was used for the protrusion of an attachment structure. In the late-mature shell, this opening became fixed in the ventral valve. Based on the posterior margin and the shell microstructure, a close relationship between Mickwitzia and the paterinids is proposed with differences interpreted as heterochronic. The shell-penetrative setal apparatus of M. muralensis is distinct from that previously described of Micrina, though both types are conceivably homologous to adult and juvenile setae of modern brachiopods.     

青海格尔木市乌丽晚二叠世乌丽群下部的腕足化石群及Waagenites的地理分布

记述的腕足动物化石标本采自青海格尔木市乌丽附近的开心岭煤矿（妈乌丽煤矿）乌丽群的下部,经研究认为,该化石群的腕足动物属种组合及生物相特征,可与华南苏皖南部、川东华蓥山的龙潭组,黔西的宣威组下部,以及藏东昌都一带的妥坝组等腕足化石群对比,属于华南晚二叠世早期（吴家坪期）海陆交互相含煤岩系的产的常见类群。讨论和纠正了英方关于该化石群的部分属种鉴定名称、时代结论,以及中国学者过去引证乌丽群化石资料与地层名称方面的错误,并就二叠纪质作用和地质历史的强烈影响,探讨了Waagenites与Tethyo-chonetes的地质地理分布。     

A relict Ordovician brachiopod fauna from the Parakidograptus acuminatus Biozone (lower Silurian) of the English Lake District

At Yewdale Beck in northern England, a brachiopod fauna dominated by species of Hindella, Kinnella, Mirorthis, Paromalomena and Plectothyrella , occurs interbedded with a graptolite fauna which includes Akidograptus ascensus , Atavograptus ceryx , Persculptograptus parvulus and Normalograptus spp. (including probable examples of N. normalis , N. angustus and N. medius ) within the lower part of the Skelgill Formation. This suggests that taxa of the terminal Ordovician Hirnantia fauna occur within the lower P. acuminatus Biozone, representing the youngest documented occurrence of the Hirnantia brachiopod fauna. Biostratigraphy, brachiopods, graptolites, Ordovician-Silurian boundary.     

Tremadoc (earliest Ordovician) brachiopods from Purmamarca and the Sierra de Mojotoro, Cordillera Oriental of northwestern Argentina

The Late Tremadoc storm-dominated shoreface to inner platform deposits exposed west of the Purmamarca village (Coquena Formation) contain a considerably more diverse brachiopod fauna than previously reported. Coquinite horizons from the lower heterolitic succession have yielded monospecific associations of Nanorthis purmamarcaensis nov. sp. (formerly assigned to N. christianiae KJERULF), which is also reported from the Late Tremadoc rocks of the Cerro San Bernardo area. The fine-grained Upper Member of the Coquena Formation contains a more diverse fauna composed by Nanorthis brachymyaria nov. sp., Astraborthis quebradensis nov. sp. and the new plectorthid genus Lipanorthis (type species L. andinus nov. sp.). A different species of Lipanorthis (L. santalaurae nov. sp.) from the Mid Tremadoc Floresta Formation of the Sierra de Mojotoro is also described.     

Intermediary metabolism and shell growth in the brachiopod Terebratalia transversa

Juvenile Terebratalia transversa (Brachiopoda) metabolize carbohydrates in the anterior-most marginal mantle at a rate of 0.46 μM glucose/g/hr (in vitro incubation of mantle in C¹⁴-glucose in a carrying medium of 10^-3 M non-radioactive glucose). The rate declines to 0.18μM glucose/g/hr in full-grown specimens. Carbohydrate metabolism in the marginal (anterior-most) mantle averages approximately 3.7 times greater than metabolism in (a portion of the ‘posterior’) mantle situated between the coelomic canals and the marginal mantle. This ratio remains constant in specimens of all sizes (i.e. an ontogenetic trend in the ratio is absent at p≤ 0.05). Organic acids are not detectable within the mantle (HPLC techniques) even after simulated anoxia (N₂ bubbling during mantle incubation). Glucose metabolism in vitro declines in both the marginal and ‘posterior’ mantles during anoxia and the metabolic ratio between marginal/‘posterior’ mantles becomes 1/1. We found no difference (at p≤ 0.05) in mean metabolic activity or in sue-related metabolic trends among populations from depths ranging between mean sea level and 70 m. However, the activity within the ‘posterior’ mantle was more variable in specimens from 70 m than in those from shallower habitats (10 m - mean sea level). The size of the specimens analyzed was most variable in the groups obtained from the shallowest habitats and least variable at 70 m depth. Our results may help define the energetics of fossil as well as living brachiopod shell growth. Brachiopod shell growth is known to be very slow relative to that of bivalves and our results indicate that this is a result of the animals' slow metabolism. The inflation of the valves in T. transversa is, in part, a function of the high ratio of intermediary metabolism in the marginal vs‘posterior’ mantle (i.e. parallels the relative growth rates at the shell margin vs‘posterior’ areas). We found that the bivalve, Chlamys hastata, which is commonly associated with T. transversa, has a lower ratio of metabolic activities in the ventral/dorsal mantle areas than the brachiopod has in the anterior/posterior. The difference produces a flatter shell in the bivalve in accord with allometric principles. The higher metabolic rate in the marginal vs‘posterior’ brachiopod mantle and its more pronounced decline with anaerobiosis is reflected in the greater definition of growth increments in the outer shell layer. Our results do not support recent generalizations that correlate shell thickness of a wide variety of invertebrates inversely with metabolic rate. Growth rate as determined from width of shell growth increments is a better index of metabolic rate. Although the genetic basis of glucose metabolism is unknown, the observed metabolic variability is consistent with suggestions that populations of marine organisms living in stable offshore environments are genetically more variable but morphologically more uniform than populations from shallow water. Furthermore, our results support suggestions that bivalved molluscs and brachiopods are very different metabolically, but the data are neutral with respect to theories of competitive exclusion of the two taxa throughout geologic history.     

Les Rhynchonelles du Bathonien du domaineprovencal: Liaison entre l'installation des peuplements et les modalités de la transgression bathonienne dans les Alpes-Maritimes

In the Maritime Alps, the Middle and Upper Bathoniancorresponds to a transgressive episode on the Provençal platform. The advancement of the marine invasion is very noticeable between Grasse and Antibes. The sediment shows the existence of a shallow marine environment in which benthic organisms were predominant.The deposits are at first varied: marly layers withbivalves (burrowers for the most part) and calcareous layers with monospecific populations of brachiopods (Burmirhynchia mediterranea nov. sp. in a calm environment, and B. turgidaBuckman in a more turbulent environment).The deposits then become uniform and show theexistence of very shallows environments: B. decorata kiliani nov. subsp. is the only representative of macrofauna in the micritic limestones that can be attributed to the Upper Bathonian.The Rhynchonellids, as on the other Bathonian carbonateplatforms of Western Europe, seem to be the only organisms likely to stay alive and proliferate in the specific environments in which entirely carbonate sediments are deposited in shallow water. Due to their relationships to those existing in other platforms, the species described in this article permit the clarification of biostratigraphic correlations.