Biogeochemistry of Paleozoic brachiopods from New York State and Ontario

A comprehensive elemental, isotopic and microstructural analyses was undertaken of brachiopod calcites from the Hamilton Group (Middle Devonian), Clinton Group (Middle Silurian) and Middle to Upper Ordovician strata of Ontario and New York State. The majority of specimens were microstructurally and chemically preserved in a pristine state, although a number of specimens show some degree of post-depositional alteration. Brachiopod calcites from the Hamilton and Clinton Groups were altered by marine derived waters whereas Trenton Group (Middle Ordovician) brachiopods altered in meteorically derived fluids. Analysis of the elemental and isotopic compositions of pristine Hamilton Group brachiopods indicates there are several chemical relationships inherent to brachiopod calcite. Taxonomic differentiation of Mg, Sr and Na contents was evident in three co-occuring species from the Hamilton Group. Mean Mg contents of pristine brachiopods were respectively Athyris spiriferoides (1309ppm), Mucrospirifer mucronatus (1035ppm) and Mediospirifer audacula (789ppm). Similarly, taxonomic differentiation of shell calcite compositions was observed in co-occuring brachiopods from the Clinton Group (Middle Silurian) and the Trenton Group (Middle Ordovician). The taxonomic control of elemental regulation into shell calcite is probably related to the slightly different physiological systems and secretory mechanisms. A relationship was observed in Hamilton Group species between the depth of respective brachiopod communities and their Mg, Sr and Na contents. These elements were depleted in the shell calcites of deeper brachiopods compared to their counterparts in shallower reaches. Apparently shell calcite elemental composition is related to environmental conditions of the depositional setting, which may have controlled the secretory regime, mineral morphology of shell calcite and precipitation rates of each species. Despite the change in Mg, Sr and Na contents between beds and formations in response to environmental conditions, the taxonomic differentiation of shell calcite composition is maintained. Thus, it may be possible to predict relative depth changes in paleoenvironmental reconstructions using brachiopod calcite. This relationship of brachiopod chemistry to depth was also tested within a transgressiveregressive (T-R) cycle in the Rochester Shale Formation (Middle Silurian). Decreasing Mg, Sr and Na contents were observed in the transition from the shallow carbonates of the Irondequoit Formation to the deeper shales of the lowest 2 m of Rochester Shale. However, no isotopic and elemental trends were observed within the entire T-R cycle which suggests that either the water conditions did not change significantly or that the cycle is illusory. A similar relationship was observed between the Fe and Mn chemistries of shell calcite and redox/paleo-oxygen conditions. Hamilton Group brachiopods analysed from deeper areas of the shelf are enriched in Mn and Fe relative to those from shallow zones. The presence of black shales and dysaerobic faunas, during deposition of the Hamilton Group, suggests that the waters of the northern Appalachian Basin were stratified. The deeper brachiopods were marginally positioned above an oxycline and their shell calcites reflect periodic incursions of oxygen depleted water. Furthermore, analysis of Dalmanella from the black shales of the Collingwood Shale (Upper Ordovician) in comparison to those from the carbonates of the Verulam Formation (Middle Ordovician) confirm the relationship of Fe and Mn contents to periodic but not permanent incursions of low oxygen waters. The isotopic compositions of brachiopod calcite found in Hamilton Group (813C; +2.5% 0 to +5.5% 0; 8180 -2.50/00 to -4.00/00) and Clinton Group (813C; +4.00/00 to +6.0; 8180; -1.8% 0 to -3.60/ 00) are heavier than previously reported. Uncorrected paleotemperatures (assuming normal salinity, 0% 0 SMOW and no fractionation effects) derived from these isotopic values suggest that the Clinton sea temperature (Middle Silurian) ranged from 18°C to 28°C and Hamilton seas (Middle Devonian) ranged between 24°C and 29°C. In addition, the isotopic variation of brachiopod shell calcite is significant and is related to environmental conditions. Within a single time-correlative shell bed (the Demissa Bed; Hamilton Group) a positive isotopic shift of 2-2.5% 0 in 013C compositions and a positive shift of 1.0-1.50/00 in 0180 composition of shell calcite is observed, corresponding with a deepening of brachiopod habitats toward the axis of the Appalachian Basin. Moroever, a faunal succession from deeper Ambocoelia dominated brachiopod association to a shallow Tropidoleptus dominated assocation is reflected by isotopic shifts of 1.0-1.50/00. Although, other studies have emphasized the significance of ±20/oo shifts in brachiopod isotopic compositions, the recognition of isotopic variability in brachiopod calcite within single beds and within depositional settings such as the Appalachian Basin has important implications for the interpretation of secular isotopic trends. A significant proportion of the variation observed isotopic distribution during the Paleozoic is related to environmental conditions within the depositional setting.

Diagenesis and geochemical stratigraphy of Upper Pennsylvanian madera brachiopods, New Mexico

Owing to the fact that low-Mg calcite fossil shells are so important in paleoceanographic research, 249 brachiopod, cement and matrix specimens from two neighboring localities (Jemez Springs and Battleship Rock), of the Upper Pennsylvanian Madera Formation were analyzed. Of which, about 86% of the Madera brachiopods are preserved in their pristine mineralogy, microstructure and geochemistry. Cement and matrix samples, in contrast, have been subjected to complete but variable post-deposition~1 alteration. It is confirmed that the stable isotope data of brachiopods are much better than that of matrix material in defining depositional parameters. Because there is no uniform or constant relationship between the two data bases (e.g., from 0.1 to 3.0%0 for 0180 and from 0.2 to 6.7%0 for 013C in this study), it is not possible to make corrections for the matrix data. Regarding the two stratigraphic sections, elemental and petrographic analyses suggest that Jemez Springs is closer to Penasco Uplift than Battleship Rock. Seawater at Jemez Springs is more aerobic, and the water chemistry is more influenced by continental sources than that at Battleship Rock. In addition, there is a relatively stronger dolomitization in the mid-section of the Battleship Rock. Results further suggest that no significant biogenic fractionation or vital effects occurred during their shell secretion, suggesting that the Madera brachiopods incorporated oxygen and carbon isotopes in equilibrium with the ambient seawater. This conclusion is not only drawn from the temporal and spatial analyses, but also supported by brachiopod inter-generic comparison (Composita and Neospirifer) and statistical analysis ( t-test).

Middle permian brachiopods from central Peninsular Malaysia - faunal affinities between Malaysia and west Cambodia

A moderately diverse Permian brachiopod fauna is described from a new rock unit, the Bera Formation, in the Bera District, central Pahang, Peninsular Malaysia. The fauna consists of 19 taxa, including 14 genera and 17 (both identified and unidentified) typically Tethyan species. The fauna appears to be correlative on the basis of brachiopods with the Neoschwagerina-Yabeina fusulinid Zones in Indochina and South China. In particular, it has strong linkage to Member C (Yabeina beds) of the Sisophon Limestone, west Cambodia. This is indicated by three of the Bera species — Urushtenoidea chaoi (Ching), Spyridiophora gubleri Termier and Termier, and Transennatia termierorum sp. nov., being shared with the Cambodian fauna. A possible early Capitanian (Middle Permian) age is proposed for the Bera brachiopod fauna.

Permian brachiopods from the Selong Xishan section, Xizang (Tibet), China. Part 2: palaeobiogeographical and palaeoecological implications, spiriferida, athyridida and terebratulida

Brachiopods from the Selong Group at the Selong Xishan section indicate a typical Peri-Gondwanan affinity, but possibly a rapid invasion of tropical/subtropical Tethyan elements into the Himalayan region at the very end of Permian. A brief analysis of taphonomy and composition of the Selong fauna indicate that Selong Xishan lay on the shallow continental shelf near shore and brachiopods suffered substantial abrasion from contemporaneous wave action. Species of Spiriferida, Athyridida and Terebratulida are herein described and revised. New taxa are Nakmusiella selongensis nov. gen. et sp. and ? Bullarina striata nov. sp.

Middle permian (Guadalupian) brachiopods from the Xiujimqinqi area, Inner Mongolia, northeast China, and their palaeobiogeographical and palaeogeographical significance

A small brachiopod fauna is described from the lower part of the Xiujimqinqi Formation of the Xiujimqinqi area in central-east Inner Mongolia, northeast China. The age of this fauna is regarded as Wordian (Middle Guadalupian, Middle Permian) by comparison with a similar brachiopod fauna from the Zhesi area of central Inner Mongolia, and by constraints from fusulinaceans associated with the Zhesi fauna. The Xiujimqinqi fauna is typical of mixed Boreal/Palaeoequatorial Middle Permian brachiopod faunas of East Asia. The mixed nature of these faunas is interpreted to have resulted from the combined effects of a middle palaeolatitudinal position, intensified plate convergence between Sino-Korea and Mongolia, and sea surface current connections with both the Arctic Sea in the north and eastern Palaeo-Tethys to the south. Possible Kaninospirifer is reported for the first time from China.

Early carboniferous athyridid brachiopods from the Qaidam Basin, Northwest China

Four new Early Carboniferous athyridid species in three genera, including one new genus, Bruntonathyris, are described from the Qaidam Basin, northwest China: Lamellosathyris qaidamensis, Bruntonathyris amunikeensis, Bruntonathyris? heijianshanensis, and Lochengia qinghaiensis. Based on the new material and also on published information, we also reviewed the taxonomic composition and the stratigraphic and paleogeographic distributions of the three genera. As a result, Lamellosathyris is considered to be indicative of late Famennian to Viséan age, originating in late Famennian in central North America and Armenia of Russia, respectively. Later, the genus appears to have two migratory directions: one branch rapidly dispersed over Mississippi Valley, Oklahoma, Texas and New Mexico of central North America in Tournaisian; alternatively, another branch from Armenia migrated westerly to Belgium, France, Spain, Britain, Ireland, via the Moscow Basin and Ural seaway, eastward to the Tienshan Mountains and Qaidam Basin of northwest China during the Tournaisian to Viséan, and easterly along the southern shelves of the Paleo-Tethys to Iran and western Yunnan of southwestern China in Tournaisian. Both Bruntonathyris and Lochengia are restrictedly Tournaisian to Viséan in age, and probably originated in the Qaidam Basin. Later, Bruntonathyris migrated easterly to South China and Japan, and westerly to Urals, Moscow Basin, Donetsk Basin and Britain; Lochengia migrated easterly to South China and westerly to the Urals seaway and the adjoined Russian Platform (i.e., both the Moscow and Donetsk Basins).

Guadalupian (Middle Permian) brachiopods from Sungai Toh, a Leptodus Shale locality in the Central Belt of Peninsular Malaysia. Part I: Lower Horizons

This work presents a systematic study of Permian Brachiopoda from the Sungai Toh Leptodus Shale locality, Pahang State, Peninsular Malaysia. This locality lies within the Central Belt of Peninsular Malaysia, a tectonic unit characterised.by tuffaceous sediments and limestones of Late Palaeozoic age. Two brachiopod-bearing horizons were studied in detail at this locality, the lower one (Horizon 2) bearing a mixed plant and invertebrate assemblage, including the brachiopods Urushtenoidea chaoi (CHING), Leptodus richthofeni KAYSER, Anidanthus cf. sinosus HUANG, Acosarina dorashamensis (SOKOLSKAJA), A. minuta (ABleH) and unidentifiable species of Linoproduetus, Neochonetes, and Strophalosiina. Horizon 3 contains a more abundant and diverse brachiopod fauna, comprising a total. of 57 species representing 47 genera, including Vediproductus punetatiformis (CHAO), Permianella typica HE & ZHU, Tranrennatia gratiosa (WAAGEN), Leptodus richthofeni KAYSER, Leptodus cf. tenuis (WAAGEN) and "Semibrachythyrina" [= Alphaneospirifer] cf. pyramidiformis LIANG. It is
suggested in this study that the age of the Sungai Toh locality is Capitanian (late Guadalupian) to possibly Wuchiapingian (early Lopingian),
as it appears to correlate well with the Lengwu fauna from Zhejiang in eastern China. The palaeobiogeographical affinities of the Sungai Toh fauna are interesting, mainly indicating strong Palaeo-equatorial affinities, while there are also some elements more typical of the cooler periGondwana
Region.

Paleobiogeographical extinction patterns of Permian brachiopods in the Asian-western Pacific region

Spatial and temporal variations in biological diversity are critical in understanding the role of biogeographical regulation (if any) on mass extinctions. An analysis based on a latest database of the stratigraphic ranges of 89 Permian brachiopod families, 422 genera, and 2059 species within the Boreal, Paleoequatorial, and Gondwanan Realms in the Asian–western Pacific region suggests two discrete mass extinctions, each possibly with different causes. Using species/family rarefaction analysis, we constructed diversity curves for late Artinskian–Kungurian, Roadian–Wordian, Capitanian, and Wuchiapingian intervals for filtering out uneven sampling intensities. The end-Changhsingian (latest Permian) extinction eliminated 87–90% of genera and 94–96% of species of Brachiopoda. The timing of the end-Changhsingian extinction of brachiopods in the carbonate settings of South China and southern Tibet indicates that brachiopods suffered a rapid extinction within a short interval just below the Permian/Triassic boundary.

In comparison, the end-Guadalupian/late Guadalupian extinction is less profound and varies temporally in different realms. Brachiopods in the western Pacific sector of the Boreal Realm nearly disappeared by the end-Guadalupian but experienced a relatively long-term press extinction spanning the entire Guadalupian in the Gondwanan Realm. The end-Guadalupian brachiopod diversity fall is not well reflected at the timescale used here in the Paleoequatorial Realm because the life-depleted early Wuchiapingian was overlapped by a rapid radiation phase in the late Wuchiapingian. The Guadalupian fall appears to be related to the dramatic reduction of habitat area for the brachiopods, which itself is associated with the withdrawal of seawater from continental Pangea and the closure of the Sino-Mongolian seaway by the and-Guadalupian.

Middle Permian brachiopods from the Dongujimqinqi area, Inner Mongolia, China

A small collection of brachiopods is described from the lower Middle Permian Yanchibeishan Formation of the Dongujimqinqi area, eastern Inner Mongolia, China. This fauna consists of the following four species: Kochiproductus sp., Linoproductus simensis (Tschernyschew), Rhynchopora inconstantis Lee and Gu and Licharewia grewingki (Netschajew). The Dongujimqinqi fauna exhibits a strong Boreal-type aspect and suggests an early Middle Permian in age.

Permian brachiopods from the Baoshan and Simao Blocks in Western Yunnan, China

This paper describes and illustrates four Permian brachiopod assemblages (Assemblages A–D) from western Yunnan, southwest China. Assemblage A occurs in the basal part of the Yongde Formation in the Xiaoxinzhai section in the southern Baoshan Block and is assignable to the latest Artinskian or earliest Kungurian. Assemblage B occurs about 50 m stratigraphically above Assemblage A in the same section, and is of Chihsian (Kungurian–Roadian) age. Assemblage C was collected from the Yongde Formation in the Anpaitian section in the southern Baoshan Block and is most likely Roadian to Wordian in age. Assemblage D came from the Longtan Formation in the Simao Block and is of late Wuchiapingian in age. The brachiopod faunas from the Yongde Formation of the Baoshan Block, as a whole, is dominated by species characteristic of the Cathaysian Province, although some links with Peri-Gondwanan faunas are also evident. In contrast, Assemblage D from the Simao Block is characterised exclusively by taxa of the Cathaysian Province. New taxa described and illustrated are Neochonetes (Huangichonetes) inflatus n. sp. Costatumulus minor n. sp. and Celebetus yunnanensis n. sp.

Permian brachiopods from the Tupe formation, San Juan Province, Precordillera, Argentina

The brachiopod fauna from the Tupe Formation at La Herradura Creek, located on the west flank of Perico Hill, San Juan Province, Argentina, palaeogeographically belongs to the western sector of the Paganzo basin ('Guandacol embayment'). The stratigraphical section of the Tupe Formation at La Herradura Creek is the stratotype of the Tivertonia jachalensis-Streptorhynchus inaequiornatus biozone, was previously regarded as being of Late Carboniferous age but here is assigned to the earliest Permian (Asselian). We describe and review the biozone assemblage, which consists of Streptorhynchus inaequiornatus, Tivertonia jachalensis, Kochiproductus sp., Costatumulus sp., Coronalosia argentinensis, Tupelosia paganzoensis, Trigonotreta pericoensis, Septosyringothyris sp. aff. Septosyringothyris jaguelensis and Crurithyris? sp. This brachiopod assemblage is related to Indian and Australian Early Permian faunas and its presence in the La Herradura Creek section provides new evidence in support of an Asselian (Early Permian) age for the Tivertonia jachalensis-Streptorhynchus inaequiornatus biozone. This assemblage is also important for intra- and inter-basinal correlation because several of its characteristic species have been identified from other sections of the Paganzo basin and the Riacuteo Blanco basin. The proposed age for this biozone is consistent with the age of palynological data from slightly above the marine faunas from the stratotype locality.