From bivalves to birds : oxidative stress and longevity

1. The oxidative stress theory of ageing predicts that animals living longer will have less cumulative oxidative damage together with structural characteristics that make them more resistant to oxidative damage itself.
2. Although a general relationship between body size, metabolism and longevity does not exist in marine invertebrates, they are generally characterized by low rates of metabolism and reactive oxygen species (ROS) formation associated with lower antioxidant enzyme activities compared to vertebrates.
3. Birds and mammals have very similar size-affected metabolic rates and their metabolic intensity explains only some of the variation in maximum lifespan potential (MLSP).Within each class, smaller animals have higher rates of metabolism and ROS production and membranes that are more susceptible to oxidative damage and autocatalytic propagation of free radicals than larger ones.
4. Although the high variation in life-history strategies is accompanied by substantial variation in MLSP, there is a consistent positive correlation between rates of ROS formation and antioxidant levels among most animals examined so far for these traits. The consensus of these studies is that ROS and antioxidant levels are inversely related to MLSP.
5. The lack of a clear stoichiometric relation between variables contributing to oxidative stress limits our capacity to infer longevity consequences from measures of pro-oxidant or antioxidant status among or within species

Distributional overlap rather than habitat differentiation characterizes co-occurrence of bivalves in intertidal soft sediment systems

Diverse species assemblages are often associated with a diversity of habitat structures. Sedimentary systems seem to be no exception, as within sedimentary systems benthic species diversity within a sample point appears to correlate with sediment grain size complexity. However, it remains to be shown whether total benthic species diversity relates to a system’s sediment heterogeneity across multiple systems. In the present paper we examined whether bivalve diversity is associated with: (1) sediment heterogeneity across systems and (2) sediment grain size complexity within systems, at 9 temperate and tropical tidal flat systems. Although bivalve life-history strategies, like post-settlement habitat selection, might suggest that sediment heterogeneity should be important for bivalve species, bivalve diversity and sediment heterogeneity were not associated across systems. Interestingly, the association between total benthic diversity and sediment heterogeneity was also not significant, suggesting that changing species composition across systems does not account for the lack of a correlation between bivalve diversity and sediment heterogeneity. Instead of habitat differentiation, bivalve diversity within a sample point was highest in ‘complex’ fine-grained sediments and bivalve distributions showed a large degree of distributional overlap in all systems. The results of this study at both smaller and larger spatial scales suggest that coexistence between bivalve species in diverse tidal flats is not associated with increased sediment heterogeneity.

Repeatable sediment associations of burrowing bivalves across six European tidal flat systems

Burrowing bivalves are associated with particular sediment types within sedimentary systems. The degree to which bivalve sediment associations are repeatable across systems has seldom been investigated. To investigate whether such repeatability exists across tidal flats, we compared adult and juvenile distributions of 3 bivalve species (Cerastoderma edule, Scrobicularia plana, Macoma balthica) across 6 European tidal flats. Across systems, the adult bivalves showed fairly repeatable distributions, with C. edule occurring in sandy sediments and M. balthica and S. plana occurring in muddy sediments. Exceptions were observed in systems composed primarily of muddy sediments (Aiguillon Bay and Marennes-Oléron Bay) and the Dutch Wadden Sea. Interestingly, juveniles and adults of C. edule and S. plana showed similar distributions across systems. M. balthica juveniles and adults showed habitat separation in 3 of the 6 studied systems; in 2 of these, it has been shown previously that juvenile M. balthica settle in mud at high tidal levels and migrate to lower sandier flats later in life. The high occurrence of juvenile M. balthica towards high sandy flats in Mont Saint-Michel Bay suggests that juveniles might choose high tidal flats rather than muddy sediments per se. A repeatable association in adults and juveniles with respect to sediment could suggest that juveniles actively settle in the proximity of the adults and/or that juveniles settling away from the adults incur a higher mortality due to either predation, physiological stress, or other factors.

How small-scale variation in oyster reef patchiness influences predation on bivalves

As oyster fishing continues to degrade reef habitat along the US Atlantic coast, oyster reefs appear increasingly fragmented on small spatial scales. In outdoor mesocosms, experiments tested how consumption of representatives of 4 different bivalve guilds by each of 3 mesopredators varies between continuous and fine-scale patches of oyster reef habitat. The mesopredator that fed least (stone crab) exhibited no detectable change in consumption on any bivalve (ribbed mussel, bay scallop, hard clam, and 3 size classes of eastern oyster). Consumption of bay scallops by both blue crabs and sheepshead fish was greater in small patches than in continuous oyster reef habitat. Of the bivalve guilds tested, only the scallop possesses swimming motility sufficient to reduce predation, an escape response that would likely leave the bivalve protected within structured habitat in larger continuous oyster reefs. Sheepshead consumed more small oysters in the continuous habitat than in the fine patches, while no other predator-prey interaction exhibited differential feeding as a function of habitat patchiness. Consequently, predation by mesopredators on bivalves can vary with the scale of oyster reef patchiness, but this process may depend upon the bivalve guild. Understanding the role of habitat patchiness on fine scales may be increasingly important in view of the declines in apex predatory sharks leading to mesopredator release, and global climate change directly and indirectly enhancing stone crab abundances, thereby increasing potential predation on bivalves.

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.

Growth rates of the infaunal bivalve Soletellina alba (Lamarck, 1818) (Bivalvia: Psammobiidae) in an intermittent estuary of southern Australia

Caging and a mark–recapture design were used to estimate the growth rate of the brittle, infaunal bivalve Soletellina alba in the Hopkins River estuary. The growth of both caged and uncaged individuals was monitored at three sites near the mouth of the estuary over 180 days. Growth rates did not differ for caged and uncaged bivalves, or for bivalves subject to different amounts of handling, or between sites. Growth did differ between consecutive time intervals, which was attributable to negligible growth occurring during the colder months of autumn/winter. Comparisons of the condition (as indicated by total mass for length³) of S. alba were inconsistent between sites for caged and uncaged bivalves and for those subject to different amounts of handling. Soletellina alba is a rapidly growing bivalve with mean growth rates for the three time intervals being 0.04±0.002 mm day⁻¹ in summer, 0.02±0.001 mm day⁻¹ in autumn and 0.03±0.001 mm day⁻¹ from summer to winter. Using existing literature, it was shown that a significant relationship exists between maximum shell length and onset of sexual maturity in bivalve molluscs. This relationship predicts that S. alba should reach the onset of sexual maturity at 15.8 mm length. Therefore, it appears that it may be possible for juvenile S. alba (<1 mm) to grow, reach sexual maturity and reproduce in between annual mass-mortality events caused by winter flooding.

Effect of flooding on estuarine bivalve populations near the mouth of the Hopkins River, Victoria, Australia

The densities of two common intertidal/shallow subtidal bivalves, Soletellina alba and Arthritica helmsi, were sampled in vegetated and unvegetated habitats of the Hopkins River estuary on three occasions during the  autumn/winter 1995.Winter flooding coincided with mass mortalities of the infaunal bivalve S. alba, but not A. helmsi. Mortalities were apparent for individuals living deeper in the sediment (≈35 cm) in vegetated and unvegetated habitats, but small S. alba (<1mm) were less susceptible to mortality than larger individuals (>1mm). Mortalities were similar across different habitat types and sediment depths, and at multiple sites within close proximity to the estuary mouth.

Effect of lowered salinity on the survival, condition and reburial of Soletellina alba (Lamarck, 1818) (Bivalvia: Psammobiidae)

Mass mortalities of fauna are known to occur in estuarine environments during flood events. Specific factors associated with these mortalities have rarely been examined. Therefore, the effect of exposing, to lowered salinities, an infaunal bivalve that is susceptible to mass mortalities during winter flooding in a southern Australian estuary was tested in the present study. In a laboratory experiment, low salinities (≤6 parts per thousand [ppt]), which mimicked those expected during flood events in the Hopkins River estuary, were shown to affect Soletellina alba, both lethally and sublethally. All bivalves died at 1 ppt, while those at 6 ppt took longer to burrow and exhibited a poorer condition than those at 14 and 27 ppt. The limited salinity tolerance of S. alba suggests that lowered salinities are a likely cause of mass mortality for this species during winter flooding.

New ophiuroids from the Permian/Triassic boundary beds of South China

A new genus of ophiuroid, Huangzhishania, is created based on new material from the Permian/Triassic boundary beds at the Huangzhishan section, South China. The age of the new genus is constrained as earliest Griesbachian by means of faunal correlation of the associated bivalves and stratigraphical correlation with the Mixed Fauna Beds of the neighbouring Meishan section. Taphonomic and palaeoecological evidence suggest that the collapse of the ophiuroid association was related to a catastrophic event, and Huangzhishania was rapidly buried in life position.

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.

Carboniferous brachiopods from the "Levipustula Fauna" in central-western Argentina : biostratigraphic, palaeoclimatic, and palaeobiogeographic implications

The “Levipustula fauna” is a relatively diverse fossil assemblage composed of brachiopods, bivalves, bryozoans, gastropods and crinoids that appears in glaciomarine sequences related to the Carboniferous glacial event that affected the central-western Argentinean basins. Brachiopods that characterize this fauna have been studied in some of the classical localities of the Argentine Precordillera in San Juan province.

In the Hoyada Verde Formation, the “Levipustula fauna” is usually located immediately above the glacial diamictite horizons and appears to be associated with the mudstones facies of the postglacial transgression. From a palaeoecological study, three brachiopod subfaunas are distinguished in this formation: the lower “Neospirifer” (“Trigonotretidae” gen. nov.) subfauna, above which the more diversified Kitakamithyris subfauna occurs, and the upper Levipustula subfauna, with the dominant species being attributed to Levipustula levis Maxwell. The Hoyada Verde fauna, as well those identified in the La Capilla Formation (Cerro El Morado locality), have been proposed as the typical “Levipustula fauna”. However, in the Leoncito Formation, the “Levipustula fauna” occurs in sandstones horizons located below the glacial diamictite beds. This fauna is poorly diversified and the brachiopods are characterized by “Neospirifer” (“Trigonotretidae” gen. nov.)-Septosyringothyris assemblage and Levipustula is not abundant. This fauna has been considered a colder “pre-interglacial fauna”.

The significant taxonomic, palaeoecologic and taphonomic variations of the “Levipustula fauna”, as well as its position in the glacial sequences, suggest an important palaeoenvironmental control related to Carboniferous glacial dynamics. From the palaeobiogeographic viewpoint, this fauna shows the highest affinity with the Eastern Australian basins from where it was previously described.

In situ growth of Soletellina alba (Bivalvia: Psammobiidae) in response to detrital supply and mouth status in a seasonally-closed estuary

The supply of detritus is an important food source for many soft-sediment invertebrates, but its importance for their growth and condition is rarely, if ever, tested directly using manipulative field experiments. Therefore, we designed such a study to: (1) test the importance of fine particulate organic matter for the growth and condition of the infaunal bivalve Soletellina alba; (2) indirectly test the feeding mode of S. alba, which has been assumed to be a deposit feeder like other members of the same superfamily (Tellinoidea); (3) compare growth rates across two summers with contrasting patterns of estuary mouth opening/closing; and (4) compare the condition of individuals used in two field studies (i.e. present versus past) and a past laboratory study. Neither growth nor condition differed when organic content of the sediments was varied, which suggests that S. alba is either a suspension feeder or capable of switching modes of feeding. There was considerable interannual variation in growth with greater growth occurring during the summer with a longer period of mouth opening. This suggests that periods of mouth closure may reduce secondary production within seasonally-closed estuaries. Potential artefacts associated with laboratory trials were also identified, with laboratory bivalves exhibiting poorer condition than those used in two field trials. The present study provides no evidence that variable quantities and qualities of organic matter within the sediments influence the growth and condition of S. alba, but future studies should focus on food supplied via the water column when the estuary is open versus closed.

Population dynamics of the infaunal bivalve, Soletellina alba

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.

A late Changhsingian (latest Permian) deep-water brachiopod fauna from Guizhou, South China

A deep-water brachiopod fauna (20 species in 19 genera) is described from the Late Permian Shaiwa Group of Ziyun, Guizhou, South China. New species include Pygmochonetes? shaiwaensis and Martinia ziyunensis. This fauna is associated with deep-water assemblages of pelagic radiolarians, foraminifers, bivalves and ammonoids. The brachiopod faunal correlations and age constraints of the associated fossil groups suggest that the Shaiwa fauna is late Changhsingian (latest Permian) in age. The Shaiwa fauna superficially resembles the coeval deep-water assemblage from Guangxi, South China; both are characterized by a mixture of deep-water brachiopods and shallow-water elements.