A new genus of Rhynchonellid Brachiopod from the Lower Triassic of South China and Implications for timing the recovery of Brachiopoda after the end-Permian mass extinction

A new genus, Meishanorhynchia, is proposed based on new material from the Lower Triassic of the Meishan section, South China. It is of a late Griesbachian age based on both associated biozones (ammonoids and
bivalves) and radiometric dates of the intercalated volcanic ash beds. Comparison with both Palaeozoic and Mesozoic-Cenozoic-related genera suggests that it may represent the first radiation of progenitor brachiopods in the aftermath of the end-Permian extinction. The lowest brachiopod horizon that contains the genus is estimated to be about 250.1±0.3 Ma. This implies that the initial stage of recovery of Brachiopoda in the Early Triassic was probably about 1.3±0.3 myr after the major pulse of the end-Permian mass extinction (dated as 251.4±0.3 Ma). This is in agreement with Hallam's expectancy that biotic recovery typically begins within one million years or so of major mass extinctions, in contrast to current views on the end-Permian extinction event which propose that the recovery of most if not all biotic groups in the Early Triassic was severely delayed and only began about five million years after the end-Permian extinction.

Systematics and palaeoecology of Changhsingian (Late Permian) Ambocoeliidae brachiopods from South China and implications for the end-Permian mass extinction

Four Ambocoeliidae brachiopod species including one new species (Crurithyris tazawai sp. nov., Crurithyris sp., Paracrurithyris pygmaea and Attenuatella mengi) are described from the Changhsingian (Late Permian) deep-water facies of South China. Analysis of the morphology, palaeoecology and palaeogeographical and temporal distributions of these species revealed that the presence of a delthyrium and/or the micro-ornaments among three of the four species (Crurithyris tazawai sp. nov., Paracrurithyris pygmaea and Attenuatella mengi) favoured an epifaunal (epiphytic) lifestyle. Morphological differences suggest that Paracrurithyris pygmaea may have been more effective metabolically in forming the shell compared with Attenuatella mengi and Crurithyris tazawai. The temporal and palaeogeographical distribution of Attenuatella suggests that A. mengi inhabited cool or cold deep waters. Both Crurithyris tazawai and Attenuatella mengi disappeared earlier in the stratigraphic record than Paracrurithyis pygmaea during the Permian–Triassic mass extinction. These differences in timing of extinction, morphology and palaeogeographical distributions suggest that oxygen deficiency and trophic resource limitation (a consequence of the changing composition of marine phytoplankton in the seas) may have contributed to the end-Permian mass extinction.

Significant pre-mass extinction animal body-size changes : evidences from the Permian–Triassic boundary brachiopod faunas of South China

This paper has undertaken a quantitative and statistical analysis of brachiopod body-size changes through the marine Permian–Triassic boundary section at Zhongzhai, Guizhou Province, South China, and found that (1) pre-mass extinction dwarfing is evident for at least the rugosochonetid species chosen for this study; (2) Tethyochonetes species reduced their size earlier than that in the Neochonetes species; and (3) no significant size reduction occurred in the newly evolved species of these two genera. Inter-species competition for resources between Neochonetes species and Tethyochonetes species and the reduction of food supply in the upper part of the uppermost Permian is here proposed to explain these observed stratigraphic patterns of brachiopod body-size changes throughout the Zhongzhai section. In the case of the newly evolved species showing no significant body-size change, morphological innovations (adaptations) in the process of speciation are considered to have significantly enhanced these newly evolved species' flexibility and survival in coping with degrading environmental conditions.

Sedimentology and ichnology of two lower Triassic sections in South China: implications for the biotic recovery following the end-Permian mass extinction

Biotic recovery following the end-Permian mass extinction was investigated using trace fossil and facies analysis of two Lower–Middle Triassic sections in South China. The Susong section (Lower Yangtze Sedimentary Province) comprises a range of carbonate and mudstone facies that record overall shallowing from offshore to intertidal settings. The Tianshengqiao section (Upper Yangtze Sedimentary Province) consists of mixed carbonate and siliciclastic facies deposited in shallow marine to offshore settings. Griesbachian to Dienerian ichnological records in both sections are characterized by low ichnodiversity, low ichnofabric indices (1–2) and low bedding plane bioturbation indices (1–2). Higher ichnofabric indices (3 and 4), corresponding to a dense population of diminutive ichnotaxon, in the Tianshengqiao section suggest opportunistic infaunal biotic activity during the earliest Triassic. Ichnological data from the Susong section show an increase in ichnodiversity during the late Smithian with 11 ichnogenera identified and increased ichnofabric indices of 4–5 and bedding plane bioturbation indices of 3–5. Although complex traces such as Rhizocorallium are present in Spathian-aged strata in this section, low ichnodiversity and ichnofabric indices and diminutive Planolites suggest a decline in recovery. In the Tianshengqiao section, ichnofabric indices are moderate to high (3–5) although only six ichnogenera are present and Planolites burrows are consistently small in Smithian and Spathian strata. Complex traces, such as large Rhizocorallium and Thalassinoides, and large Planolites, did not appear until the Anisian. Ichnological results from both sections record the response of organisms to unfavourable environmental conditions although the Susong section shows earlier recovery during the Smithian prior to latest Smithian–Spathian decline. This decline may have resulted from a resurgence of euxinic to anoxic marine environment in various regions of South China. Ichnological data from the Tianshengqiao section indicate protracted recovery throughout the Early Triassic as previously found elsewhere in South China. Comparison of the South China trace fossil records with global ichnological data show a diachronous pattern of recovery of trace makers and highlights the heterogeneous development of oxic facies on the marked variation in recovery rate.

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 that Permophricodothyris, 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 as Squamularia in fact belong to Permophricodothyris. The new data presented in this paper also allows a critical evaluation of Permophricodothyris in relation to its closest allies: Phricodothyris, Squamularia, Bullarina and Neophricodothyris. The revision reveals that a total of 18 Permophricodothyris 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) and P. guangxiensis Han, Zhou & Wang, are redescribed here, providing critical new information on the morphology and taxonomy of these species.

Some morphometric and biochemical features of ready-to-migrate silver and pre-migratory yellow stages of the shortfin eel of south-eastern Australian waters

The mean total length (L_T), mass and age of ready to migrate female silver shortfin eels Anguilla australis from the Hopkins River estuary and the mouth of the Merri River in south-eastern Australia, were 83·2 ± 1·2 cm, 1051 ± 51 g, and 17·2 ± 1·79 years, respectively. The eye index (I_E) of the silver shortfin eels was < 5·2 (mean 7·64 ± 0·29) and differed significantly from that of the yellow shortfin eels collected from two other sites. The I_E increased with L_T (mm) and was related by log I_E= 2·656 log L_T6·925. The per cent moisture, protein and ash content of the liver of silver shortfin eels was significantly lower than in yellow shortfin eels, but lipid content was significantly higher in the former (35·5 ± 2·0%). The mean mass μg mg lipid ‾) of saturates (230·4 ± 2·6 v. 181·7 ±2·6), monoenes (367·4 ± 6·3 v. 290·8 ± 8·9) and PUFA (177·3 ± 5·3 v. 159·7 ± 4·6) in muscle was significantly higher, and the great majority of individual fatty acids was found also in higher quantities in silver shortfin eels. In the liver, the PUFA found in the highest quantity was 22:6n-3, except in shortfin eels from Hopkins River estuary, and the amount of 18:2n-6 in the liver of silver shortfin eels was significantly higher than that in yellow shortfin eels but the reverse was true of 20:4n-6. In both muscle and liver tissues the saturate 16:0 and the monoene 18:ln-9 collectively accounted for >50% of all the fatty acids in the lipid.

Significance of Claraia from the late permian of South Guizhou, China

Recently collecte material of two Claraia taxa, Claraia zhiyunica Yang et al, 2001 and Claraia sp. nov. from the Late Permian of South China, are described. Late Permian Claraia species are compared with those from the Early Triassic, and the survival of Claraia across the mass extinction period across the Permian- Triassic boundary (PTB) is discussed.

Post-extinction brachiopod faunas from the late Permian Wuchiapingian coal series of South China

This paper describes fourteen brachiopod species in eleven genera from the Late Permian Wuchiapingian Coal Series (Lungtan Formation) of South China. Of these, the shell bed fauna from the basal Lungtan Formation is interpreted to represent the onset of the recovery of shelly faunas in the aftermath of the Guadalupian/Lopingian (G/L) mass extinction in South China. The post-extinction brachiopod faunas in the Wuchiapingian are characterized by the presence of numerous Lazarus taxa, survivors, and newly originating taxa. These elements capable of adapting their life habits were relatively more resistant to the G/L crisis. The post-extinction faunas, including survivors and the elements originating in the recovery period, have no life habit preference, but they were all adapted to a variety of newly vacated niches in the Late Permian oceans. Two new species, Meekella beipeiensis and Niutoushania chongqingensis, are described, and two Chinese genera, Niutoushania and Chengxianoproductus, are emended based on re-examination of the type specimens and new topotype materials from the Lungtan Formation.

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.

Reproductive biology of the eastern shovelnose stingaree Trygonoptera imitata from south-eastern Australia

Abstract. In applying a quantitative approach to the reproduction of Trygonoptera imitata, the present study contributes to understanding the wide diversity in the reproductive biology of the family Urolophidae and provides insights to help determine phylogenetic relationships. This localised species is taken as bycatch in several inshore fisheries and potentially impacted by a range of other anthropogenic pressures, including introduced species, particularly in shallow-water pupping areas.T. imitata can be characterised as a species of comparatively lowmatrotrophic histotrophy with an extended period of relatively large eggs in utero (5–8 months) followed by rapid growth of the embryos (4–6 months). The reproductive cycle is annual with parturition occurring during late-February–April, followed immediately by ovulation. Mean size-at-birth is ~225mm total length and there is a ~1000% gain in mean wet mass from egg (15 g) to full-term embryo in utero (150 g), the lowest reported for any viviparous batoid. Litter size increases with maternal length, reaching a maximum of seven, and sex ratio of embryos is 1 : 1. Maximum length and estimates of the maturity–ogive parameters l₅₀ and l₉₅ are similar for females and males.

Life crises on land across the Permian–Triassic boundary in South China

The western Guizhou and eastern Yunnan area of southwest China commands a unique and significant position globally in the study of Permian–Triassic boundary (PTB) events as it contains well and continuously exposed PTB sections of marine, non-marine and marginal-marine origin in the same area. By using a range of high-resolution stratigraphic methods including biostratigraphy, eventostratigraphy, chronostratigraphy and chemostratigraphy, not only are the non-marine PTB sections correlated with their marine counterparts in the study area with high-resolution, the non-marine PTB sections of the study area can also be aligned with the PTB Global Stratotype Section and Point (GSSP) at Meishan in eastern China. Plant megafossils (“megaplants”) in the study area indicate a major loss in abundance and diversity across the PTB, and no coal beds and/or seams have been found in the non-marine Lower Triassic although they are very common in the non-marine Upper Permian. The megaplants, however, did not disappear consistently across the whole area, with some elements of the Late Permian Cathaysian Gigantopteris flora surviving the PTB mass extinction and locally even extending up to the Lower Triassic. Palynomorphs exhibit a similar temporal pattern characterized by a protracted stepwise decrease from fern-dominated spores in the Late Permian to pteridosperm and gymnosperm-dominated pollen in the Early Triassic, which was however punctuated by an accelerated loss in both abundance and diversity across the PTB. Contemporaneous with the PTB crisis in the study area was the peculiar prevalence and dominance of some fungi and/or algae species.

The temporal patterns of megaplants and palynomorphs across the PTB in the study area are consistent with the regional trends of plant changes in South China, which also show a long-term decrease in species diversity from the Late Permian Wuchiapingian through the Changhsingian to the earliest Triassic, with about 48% and 77% losses of species occurring respectively in the end-Wuchiapingian and end-Changhsingian. Such consistent patterns, at both local and regional scales, contradict the hypothesis of a regional isochronous extinction of vegetation across the PTB, and hence call into question the notion that the end-Permian mass extinction was a one-hit disaster. Instead, the data from the study area and South China appears more consistent with a scenario that invokes climate change as the main driver for the observed land vegetation changes across the PTB in South China.

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.

Early Triassic lingulidae and associated marine faunas in south China

The survival strategies of Early Triassic Lingulidae fauna and its associated shallow marine faunas across the end-Permian mass extinction 250 million years ago are discussed. Three new genera and nine new species are erected. A comprehensive database of all Lingulidae species through the Late Devonian to Present is also constructed.