17 resultados para anoxic

em Deakin Research Online - Australia


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A series of field surveys were carried out on two permanent pools of the upper Glenelg River in SW Victoria, Australia. One was representative of the wider and deeper pools while the other was representative of the more-narrow and shallower pools. Both pools showed a typical seasonal cycle of warm, brackish, oxygen-poor, summer conditions and cool, oxygen-rich, low-salinity, winter conditions. The summer salinity increases were larger than expected, suggesting possible saline groundwater inflow from unidentified springs. Both pools contained anoxic water in their deeper sections but this was permanent only in the deeper pool. A simple model of the flushing rate of such anoxic pools subject to flows, such as environmental flow releases, was developed, based on an energy balance between the potential energy required to lift the anoxic layer and the kinetic energy derived from the river flow. The results were tested against and in agreement with the field measurements. The model also suggests that the anoxic layers are resilient to all but the largest environmental flows.

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The Permian of Timor in the Lesser Sunda Islands has attracted the attention of palaeontologists since the middle of the nineteenth century because of the richness, diversity and excellent state of preservation of its fauna. These abundant fossil data have been compiled and updated for the present account. The Permian rocks of Timor were deposited on the northern margin of Australia. At the present time the northern margin of Australia, in the region of Timor, is involved in a continent–arc collision, where Australia is colliding with the Banda Arcs. As a result of this collision, Permian rocks of the Australian margin have been disrupted by folding and faulting with the generation of mud-matrix mélange, and uplifted to form part of the island of Timor. Due to this tectonic disruption, it has proved difficult to establish a reliable stratigraphy for the Permian units on Timor, especially as the classic fossil collections were obtained largely from the mélange or purchased from the local people, and do not have adequate stratigraphic control. Detailed systematic, structural, stratigraphic and sedimentological studies since the 1960s have provided a firmer stratigraphic and palaeogeographic background for reconsideration of the significance of the classic fossil collections. Permian rocks on Timor belong either to a volcanic-carbonate sequence (Maubisse Formation), or to a clastic sequence (Atahoc and Cribas formations) in which volcanics are less prominent. The Permian sequences were deposited on Australian continental basement which was undergoing extension with spasmodic volcanic activity. Carbonates of the Maubisse Formation were deposited on horst blocks and volcanic edifices, while clastic sediments of the Atahoc and Cribas formations were deposited in grabens. The clastic sediments are predominantly fine-grained, derived from a distant siliciclastic source, and are interbedded with sediments derived from the volcanics and carbonates of adjacent horst blocks. Bottom conditions in the graben were often anoxic. In the present account, events on Timor during the Permian are related to the regional tectonic context, with the northward movement of Australia leading to the amelioration of the climate from sub-glacial to sub-tropical, together with the separation of crustal blocks from the northern Australian margin to form the Meso-Tethys.

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A unique marine Permian-Triassic boundary section containing rich oil source rocks has been continuously cored in a petroleum borehole from the Perth Basin of Western Australia. Such sequences, which provide a biostratigraphic and environmental record at the time of the largest extinction event of the past 500 million years, are globally rare, and this is the first to be documented in Australia. Throughout geological history there have been periods of global marine anoxia that commonly resulted in the widespread deposition of petroleum source rocks, most notably in the mid-Cretaceous and Late Jurassic. An apparent paradox is that, previously, source rocks have not been recognised in association with the Permian-Triassic boundary, despite widespread marine anoxia at this time. The Perth Basin source rocks contain abundant and unusual biomarkers, apparently related to the highly specialised and limited biota that flourished in the aftermath of the end-Permian extinction event. Local conditions may have favoured source-rock development, either due to higher productivity resulting from coastal upwelling or through enhanced preservation under strongly anoxic conditions.

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The Permian-Triassic extinction pattern in the peri-Gondwanan region is documented biostratigraphically, geochemically and sedimentologically based on three marine sequences deposited in southern Tibet and comparisons with the sections in the Salt Range, Pakistan and Kashmir. Results of biostratigraphical ranges for the marine faunas reveal an end-Permian event comparable in timing with that known at the Meishan section in low palaeolatitude as well as Spitsbergen and East Greenland in northern Boreal settings although biotic patterns earlier in the Permian vary. The previously interpreted delayed extinction (Late Griesbachian) at the Selong Xishan section is not supported by our analysis. The end-Permian event exhibits an abrupt marine faunal shift slightly beneath the Permian-Triassic boundary (PTB) from benthic taxa- to nektic taxa-dominated communities. The climate along the continental margin of Neo-Tethys was cold before the extinction event. However, a rapid climatic warming event as indicated by the southward invasion of abundant warm-water conodonts, warm-water brachiopods, calcareous sponges, and gastropods was associated with the extinction event. Stable isotopic values of δ13Ccarb, δ13Corg and δ18O show a sharp negative drop slightly before and during the extinction interval. Sedimentological and microstratigraphical analysis reveals a Late Permian regression, as marked by a Caliche Bed at the Selong Xishan section and the micaceous siltstone in the topmost part of the Qubuerga Formation at the Qubu and Tulong sections. The regression was immediately followed by a rapid transgression beneath the PTB. The basal Triassic rocks fine upward, and are dominated by dolomitic packstone/wackestone containing pyritic cubes, bioturbation and numerous tiny foraminifers, suggesting that the studied sections were deposited during the initial stage of the transgression and hence may not have been deeply affected by the anoxic event that is widely believed to characterise the zenith of the transgression.

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This study examined the factors affecting the distribution and abundance of epifaunal caridean shrimps in seagrass meadows of the Hopkins River estuary in south-western Victoria, Australia, and investigated the life history patterns of the freshwater Parana australiensis, found for the first time in estuaries. Adult and sub-adult shrimps were surveyed in seagrass meadows along the estuary over two years, and their planktonic larvae were surveyed in adjacent waters. Three species were collected. The marine Palaemon serenus occurred only near the mouth, summer to autumn, in high salinities. The marine/estuarine Macrobrachium intermedium occurred throughout the estuary. Adults were most abundant in late autumn, and least abundant in summer (unlike trends reported in marine meadows). Densities were higher and less variable in downstream meadows. P. australiensis occurred in the upper estuary all year, most abundantly in spring, due to migration from the river after peak discharge. Ovigerous females dominated, while males, showing less migration into the estuary, dominated above estuarine influence. Adults disappeared from the estuary in summer as salinity rose. Breeding period for P. australiensis was briefer in the estuary (September-December) than upstream (July-April). M. intermedium began breeding later in the upper estuary (November/December-March) than in the lower estuary (October-March), probably reflecting a physiological response to lower salinity, rather than an interaction with P. australiensis. No ovigerous P. serenus were found in the estuary. Larvae of P. australiensis and M intermedium occurred abundantly throughout the estuary, but P. serenus larvae did not. P. australiensis was an early coloniser to the plankton after peak discharge (November-December). Larvae concentrated in the deep saline layer at the head of the intruding salt wedge, thus probably maintaining longitudinal position. Diurnal vertical migrations were evident within the salt wedge, and in a deep pool above tidal influence. M. intermedium larvae occurred October-May in the lower estuary and November-April in the upper estuary, peaking in abundance one to two months after P. australiensis. They were associated with low surface flows and surface salinities greater than 10, over an anoxic deeper layer. All three species exhibited extended development of euryhaline larvae in the laboratory. Tolerances and optimal salinities of larvae of the three species reflected their distributions. M. intermedium was the most euryhaline species. P. australiensis larvae were tolerant of higher salinities than juveniles of adults: capable of developing in salinity of at least 15. Most P. australiensis juveniles recruited to the estuary November-December, after which numbers declined dramatically. After settlement, most recruits probably migrated upstream out of the estuary. Two cohorts of M. intermedium recruited to the estuary from larvae in summer (December and February), but some juveniles also migrated from adjacent coastal waters. Post-larval migration was at least as important a determinant of abundance as direct recruitment from estuarine, planktonic larvae in all three species. Distributions among seagrass meadows along the estuary were determined primarily by physico-chemical patterns driven by hydrological changes. Seasonal variations in salinity and temperature were strongly associated with seasonal variations in shrimp abundance. Salinity tolerances of adults of the three species reflected their distribution patterns. Biotic interactions were more important in determining distributions within meadows. P. australiensis, when abundant, were associated with seagrass biomass. M. intermedium were also, but when seagrass was sparsest and least extensive. The two species apparently partitioned the seagrass meadow according to depth in early summer. Laboratory experiments suggested P. australiensis was displaced from deeper water by M. intermedium. Preference for vegetative complexity and competition for position within meadows suggest the underlying importance of predation in regulating shrimp populations. A survey of south-eastern Australian estuaries found P. australiensis larvae abundant in all stable, open, well-developed, salt-wedge estuaries where adults were abundant. Adults were most abundant in low salinities among submerged leafy macrophytes. Reproductive traits of P. australiensis were compared in estuarine and fresh reaches of three rivers. Early in the breeding season, egg size was smaller, and (size-specific) egg number larger in estuaries than upstream. A trade-off between egg size and egg number resulted in no difference in total (size-specific) reproductive investment between locations. Reproductive investment tended to decrease at some locations over the breeding season, and this decrease was a result of decreased egg size in most cases. The decrease in reproductive investment probably reflected reduced food availability for the adult, while the reduced egg size was probably a response to improved conditions for larval development. In the Hopkins River, larger egg size at upstream sites was reflected in larger early stage larvae. Later stage larvae were larger in the estuary, suggesting more favourable conditions for larval development. Allozyme electrophoresis showed the P. australiensis populations in each of the three rivers to be distinct. Allozyme frequencies were not different within the Hopkins River, but upstream and estuarine locations in the Curdies and Gellibrand were different. Although some variation in reproductive traits within catchments may have been due to genotypic differences, trade-offs between egg size and number, and decreases in egg size over summer were probably due to plastic responses to environmental cues. It is proposed P. australiensis inhabits and reproduces in both estuarine and freshwater environments by plastic response to environmental conditions. Recruitment to estuaries is dependent on the presence of suitable adult, littoral habitat, and a stable salt wedge for larval retention. Estuaries are important recruitment sites for P. australiensis, potentially allowing an extra brood each year before riverine recruitment. Estuarine broods could constitute a large part of the total fecundity of P. australiensis females. Euryhaline larvae and estuarine recruitment of P. australiensis suggest marine transport of larvae between estuaries as a possible dispersal mechanism for Paratya species.

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The population dynamics of the infaunal bivalve Soletellina alba was investigated at three sites situated within close proximity to the mouth of the Hopkins River estuary. The initial study design was planned to examine the importance of winter flooding to the persistence of this bivalve mollusc within the Hopkins estuary, since mass mortalities have been observed during previous years coincident with periods of winter flooding. Unfortunately, the climatic conditions experienced during this study were atypical compared to the long-term average, so detailed sampling was limited to two, unanticipated, non-flood years rather than two, highly anticipated, flood years. This hampered my ability to conduct complete tests of the importance of winter flooding. Patterns of river discharge and the frequency and duration of mouth opening and closing differed greatly from that expected. Unexpectedly, periods of mouth closure were not always associated with periods of minimal river discharge; low salinities were another unexpected result during an extended period of mouth closure during 1998. As expected, salinities varied considerably with increasing water depth when the estuary mouth was open. Mouth closure lead to salinities becoming more uniform between water depths but hypoxic and anoxic conditions became evident via stratification in the water column at 1 m below the Australian Height Datum (AHD). Other than trends associated with increased water depth, significant variation was not evident between measurements of salinity taken from three sites within close proximity of the estuary mouth (approximately 500 m), or during changes in tide. The most pertinent anomaly was the absence of winter flooding. The distribution and abundance of juvenile and adult S. alba was variable across all Dates, Sites and Channel elevations (i.e. water depths) sampled during this study. An experimental test comparing the recruitment of juveniles at different channel elevations and in sediments of varying particle size was conducted during an exceptionally successful period of recruitment during 1999. The results of these tests showed that recruitment was greatest at the shallowest channel elevation used, and there was little evidence that sediment particle size influenced recruitment. In contrast to 1999, recruitment during 1997 or 1998 was extremely poor. Growth rates were monitored using tagged individuals held in caged and uncaged plots, which revealed that growth was highly variable among individuals, but not between Sites. These tests also revealed that growth was negligible during the colder, winter months, and that the fastest growing individuals were capable of growing 0.2 mm/day. Mixed results were obtained for tests of potential cage artifacts and the influence of handling. Caging and differing amounts of handling did not appear to influence growth, but there was evidence that cages and handling influenced bivalve condition and number of mortalities. These direct tests appeared to be the most appropriate method for determining growth rates of this species, since attempts to analyse length-frequency data were made difficult by the apparent convergence of cohorts, and shell aging is difficult due to the thin, fragile nature of the shell. As expected, mass mortalities were observed during the flood of 1996, but not during the two non-flood years of 1997 and 1998. There were, however, some considerable declines in abundances at some channel elevations during the two non-flood years. However, these declines were attributable to the complete disappearance of individuals, rather than the sudden presence of numerous, recently dead individuals that typify observed declines during winter flooding. The complete disappearance of individuals suggest that S. alba may be capable of post-settlement emigration, or that they were consumed by an unknown predator. Salinity tolerance tests showed that bivalves exposed to low salinities (≤6 ppt), exhibited poorer condition and took longer to re-burrow into sediments than those exposed to greater salinities (≥14 ppt), while death of bivalves exposed to salinities ≤1 ppt occurred after 8 days of exposure. These tests provide evidence that low salinities are probably the principal cause of mass mortalities during winter flooding, although the interaction between salinity, temperature and turbidity also deserve consideration. The results of this study indicate that certain aspects of winter flooding, especially salinity, are responsible for the mass mortalities of S. alba rather than the result of a short-lived life history. I hypothesise that the survival of very young juveniles (between 0.5 and 1 mm shell length) and rapid growth rates are important features of the life history of S. alba that explain its successful persistence within the Hopkins River estuary. The rapid rates of growth suggest that it may be possible for juveniles that survive winter flooding to grow, reach sexual maturity, and reproduce before the onset of the next flood event. Unfortunately, the increased survivorship of juveniles during periods of winter flooding was not demonstrated by this study because of the absence of winter flooding and also relatively poor recruitment. It is highly likely that this species is capable of completing it entire life cycle within the estuary since the absence of other nearby populations, together with periods of mouth closure, are likely to greatly limit the potential contribution made by larvae entering from the surrounding marine environment. This study has added considerably to our knowledge of how infauna cope with life in the intermittently closing estuaries that typify semi-arid coastlines in the Southern Hemisphere.

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A brief appraisal of marine fossils from high latitudes and episodically cold climate especially in east Australia and New Zealand during Late Palaeozoic and Early Mesozoic times shows patterns of evolution and survival that differ from those adduced for the palaeotropics and Northern Hemisphere. Examples taken from amongst phyla Scyphozoa, Bryozoa, Brachiopoda and Classes Bivalvia and Class Cephalopoda suggest these attributes:
1. Evolution and demise of species and genera proceeded at a rate close to that known for palaeotropical and Northern Hemisphere macro-invertebrates, but involved fewer families and orders.
2. Possibly, intraspecific variation was greater amongst southern palaeohemisphere Permian species than in those of the Permian palaeotropics.
3. There was no proven diminution of life at the end of the Guadalupian (Middle Permian) at southern high latitudes, where however the fossil record is meagre for this interval. Younger Wuchiapingian and Changhsingian faunas were moderately diverse.
4. There is no evidence for a high latitude Southern Hemisphere anoxic event in the Early Triassic despite claims of a world-wide anoxic interval. Nor has any substantial volcanic eruption or bolide impact left any marked traces in the sedimentary record.
5. As a consequence, some major groups such as Bryozoa and Conulariida (Staurozoa) survived the end- Permian extinction shock in the Southern Hemisphere.
6. Other major groups appear to have survived better in the south than in the north, notably, mollusc Bivalvia and Cephalopoda. It therefore appears likely that Triassic seas were restocked substantially from the Southern Hemisphere and that the Permian extinction shock was asymmetric with respect to latitudes in its distribution and affect.

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A three stage-treatment of domestic wastewater including anaerobic, anoxic and aerobic phases is employed in this study while a clarifier unit is replaced with a submerged membrane in the aerobic unit. The effects of operational parameters on the performance of a pilot scale submerged membrane bioreactor (SMBR) namely hydraulic retention time (HRT), ratio of return activated sludge (QRS), ratio of internal recycle (QIR), solid retention time (SRT) and dissolved oxygen (DO) are evaluated by simulations, using a hybrid model composed of TUDP model, oxygen transfer model, biofouling model due to extra-cellular polymeric substances (EPS) and turbulent shear model. The results showed that anaerobic HRT of 3 hours, anoxic HRT of 6 hours, QRS of 20% and QIR of 300 % are satisfactory in obtaining a high removal efficiency (>90%) of COD, NH4-N, P04-P as well as a less sludge production. An increase of sludge production causes an increase in EPS, which fouls the membrane surface and increase the cleaning cycle of membrane. Operation of 5MBR system at 2 mg/I of DO and 30 days of SRT can extend the membrane cleaning cycle dramatically. The membrane cleaning cycle however is strongly dependent on the initial and terminal specific fluxes and displays inverse power relationships to those fluxes.

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Recirculating aquaculture systems (RAS) are essential for the reduction in fresh water usage as well as the discharge of nutrients along with aquaculture effluents. A RAS consisting of an anoxic reactor, a membrane bioreactor (MBR) and a UV-disinfection unit was used to process 10,000 L/d of aquaculture effluent providing high-quality treated water for recirculation to a Barramundi fish culture. The system maintained low levels of nitrate (<20 mg/L), nitrite (<3 mg/L) and ammonia (<0.6 mg/L) in the fish tank. Permeate from the membrane that was recirculated to the fish tank contained <21 mg/L of nitrate, <2 mg/L of nitrite and 0 mg/L of ammonia. However, the rate of fouling of the membrane in the MBR was around 1.47 kPa/d, and the membrane in the MBR required cleaning due to fouling after 16 days. Cleaning of the membrane was initiated when the TMP reached around 25 to 30 kPa. In order to reduce the rate of fouling, 500 mg of powdered activated carbon (PAC) per litre of MBR volume was introduced, which decreased the rate of fouling to 0.90 kPa/d. Cleaning of membrane was needed only after 31 days of operation while maintaining the treated effluent quality. Thus the frequency of cleaning could be halved due to the introduction of PAC into the MBR.

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In South China, the Changhsingian brachiopods are extraordinarily abundant and diverse, comprising 468 species in 144 genera. However, approximately 91% of brachiopod species were eliminated during the Permian-Triassic (P-Tr) mass extinction event. Brachiopods in the aftermath of the P-Tr mass extinction were extremely rare, with only one opportunistic taxon, Lingulida, occasionally found in the Griesbachian and Smithian at a high abundance. Species-diversity of articulated brachiopods in the early Griesbachian, late Griesbachian, Dienerian, and Smithian are 35, 3, 2, and 0, respectively. Although a few of Mesozoic-type species occurred in the Griesbachian, Dienerian and Smithian, a marked diversification of brachiopods occurred in the Spathian and early Anisian and was characterised by 9 and 17 Mesozoic-type species, respectively. The diversification of brachiopods in the Spathian and early Anisian coincides with the contemporaneous expansion of the refuge zone, suggesting that the improvement of marine environmental conditions (e.g., lethally hot temperature and anoxic seawater) played a key role in brachiopod recovery after the P-Tr mass extinction.

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An enhanced membrane bioreactor (eMBR) consisting of two anoxic bioreactors (ARs) followed by an aerated membrane bioreactor (AMBR), UV-unit and a granular activated carbon (GAC) filter was employed to treat 50-100 mg/L of remazol blue BR dye. The COD of the feed was 2334 mg/L and COD:TN:TP in the feed was 119:1.87:1. A feed flow rate of 5 L/d was maintained when the dye concentration was 50 mg/L; 10 L/d of return activated sludge was recirculated to each AR from the AMBR. Once the biological system is acclimatised, 95% of dye, 99% of COD, 97% of nitrogen and 73% of phosphorus were removed at a retention time of 74.4 h. When the effluent from the AMBR was drawn at a flux rate of 6.5 L/m(2)h, the trans-membrane pressure reached 40 kPa in every 10 days. AMBR effluent was passed through the UV-unit and GAC filter to remove the dye completely.

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 B.V. Body size is a fundamental and defining character of an organism, and its variation in space and time is generally considered to be a function of its biology and interactions with its living environment. A great deal of body size related ecological and evolutionary research has been undertaken, mostly in relation to extant animals. Among the many body size-related hypotheses proposed and tested, the size-bathymetry relationship is probably the least studied. In this study, we compiled a global body size dataset of Changhsingian (Late Permian, ca. 254. Ma-252. Ma) brachiopod species from low-latitude areas (30°S-30°N) and analyzed their species diversity and body size distribution patterns in relation to the nearshore-offshore-basin bathymetric gradient. The dataset contained 1768 brachiopod specimens in 435 species referred to 159 genera and 9 orders, from 135 occurrences (localities) of 18 different palaeogeographic regions. Treating the whole of the Changhsingian Stage as a single time slice, we divided the nearshore-offshore-basin bathymetric gradient into three broad depth-related environments: nearshore, offshore and basinal environments, and compared how the species diversity and body size varied along this large-scale bathymetric gradient.Here, we report an array of complex patterns. First, we found a clear overall inverse correlation between species diversity and water depth along the nearshore-offshore-basin gradient, with most species concentrating in the nearshore environment. Second, when the median sizes of all low-latitude brachiopod species from the three environments were compared, we found that there was no significant size difference between the nearshore and offshore environments, suggesting that neither the wave base nor the hydrostatic pressure exerts a critical influence on the body size of brachiopods. On the other hand, the median sizes of brachiopods from the nearshore environment and, to a lesser extent, the offshore environment were found to be significantly larger than that of basinal brachiopods. This trend of significant size reduction in basinal brachiopods mirrors the relative low species diversity in the basinal environment, and neither can be easily explained by the tendency of decreasing food availability towards deeper sea environments. Rather, both trends are consistent with the hypothesis of an expanding Oxygen Minimum Zone (OMZ) in the bathyal (slope to deepsea) environments, where hypoxic to anoxic conditions are considered to have severely restricted the diversification of benthos and favored the relative proliferation of small-sized brachiopods. Finally, a significant difference was also found between eurybathic and stenobathic species in their body size response to the nearshore-offshore-basin gradient, in that eurybathic species (species found in all three environments) did not tend to change their body size significantly according to depth, whereas stenobathic forms (species restricted to a single environment) exhibit a decline in body size towards the basinal environment. This pattern is interpreted to suggest that bathymetrically more tolerant species are less sensitive to depth control with respect to their body size change dynamics, in contrast to stenobathic species which tend to grow larger in shallower water depths.

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Oceanic anoxia has long been considered as one of the main causes for the end-Permian mass extinction. However, the results obtained by different researchers are rather divergent from different sections, or even on the same section using the same redox proxy. This study aims to examine the causes for some of these divergent results using high-resolution pyrite framboid sampling at the Meishan GSSP section in South China. Detailed microfacies analysis shows that the uppermost Late Permian strata comprises two significantly different sedimentary facies: one characterized by silicious muddy limestone and recognized as representing autochthonous background sediments; the other distinguished by bioclastic grainstone, interpreted to be allochthonous in origin and have been transported from the nearby platform margin. These two different sedimentary facies represent two distinctly different redox conditions. Together with the facies analysis, a statistical analysis of pyrite framboids was carried out to evaluate the redox evolution across the Permian-Triassic boundary. Abundant framboids with average diameters of about 6μm are found in background sediments beneath the extinction boundary, indicating generally anoxic bottom water conditions. But this condition was punctuated by transient intervals of rapid oxygenation interpreted to have been caused by intrusion of intermittent turbidity flows. Our study also showed that anoxic conditions persisted into the immediate aftermath of the mass extinction, thereafter it was quickly followed by a relatively long period of oxic conditions (with rare framboids). However, the redox conditions returned to anoxia (with abundant pyrite framboids averaging about 5μm in diameter), accompanied by a rapid global transgression. The oxygenation manifested near the Permian-Triassic boundary coincides with the negative excursion of carbon isotope. This would imply that, contrary to previous interpretations, this great δ13C negative excursion was probably not caused by the upwelling of anoxic deep ocean waters.

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New analysis of Permian-Triassic brachiopod assemblages and body-size changes in South China provides insights into the process of the environmental crisis in the lead up to the end-Permian mass extinction. The recently discovered Daoduishan section of South China can be considered as an important auxiliary section for the study of brachiopods at the Meishan Section D of South China, the GSSP of the Permian-Triassic Boundary (PTB). This paper studied changes of the brachiopod assemblages and body sizes through the upper part of the Changxing Formation and basal Yinkeng Formation of Daoduishan. The results show that significant changes of brachiopod assemblages took place between Beds 24e and 26. Brachiopods?Prelissorhynchia sp. and Paracruirithyris pygmaea are the dominators in Beds 14-24e, while Tethyochonetes pigmaea and Paryphella spp. are the dominators in Beds 26-29. Body sizes of brachiopods significantly decreased between Beds 24e and 26 and then maintained smaller means in Beds 27-29. Studies of brachiopod morphological features indicate both Tethyochonetes and Paryphella had advantageous adaptations enabling them to copy with living in an anoxic/dysoxic and/or low-productivity environment during the Permian-Triassic crisis.