Antarctic marine species have evolved in one of the coldest and most temperature-stable marine environments on Earth. They have long been classified as being stenothermal, or having a poor capacity to resist warming. Here we show that their ability to acclimate their physiology to elevated temperatures is poor compared with species from temperate latitudes, and similar to those from the tropics. Those species that have been demonstrated to acclimate take a very long time to do so, with Antarctic fish requiring up to 21-36 days to acclimate, which is 2-4 times as long as temperate species, and invertebrates requiring between 2 and 5 months to complete wholeanimal acclimation. Investigations of upper thermal tolerance (CT max ) in Antarctic marine species have shown that as the rate of warming is reduced in experiments, CT max declines markedly, ranging from 8 to 17.5°C across 13 species at a rate of warming of 1°C day, and from 1 to 6°C at a rate of 1°C month −1 . This effect of the rate of warming on CT max also appears to be present at all latitudes. A macrophysiological analysis of long-term CT max across latitudes for marine benthic groups showed that both Antarctic and tropical species were less resistant to elevated temperatures in experiments and thus had lower warming allowances (measured as the difference between long-term CT max and experienced environmental temperature), or warming resistance, than temperate species. This makes them more at risk from warming than species from intermediate latitudes. This suggests that the variability of environmental temperature may be a major factor in dictating an organism's responses to environmental change.
Summary1. This study examined the effects of long-term culture under altered conditions on the Antarctic sea urchin, Sterechinus neumayeri. 2. Sterechinus neumayeri was cultured under the combined environmental stressors of lowered pH (À0Á3 and À0Á5 pH units) and increased temperature (+2°C) for 2 years. This time-scale covered two full reproductive cycles in this species and analyses included studies on both adult metabolism and larval development. 3. Adults took at least 6-8 months to acclimate to the altered conditions, but beyond this, there was no detectable effect of temperature or pH. 4. Animals were spawned after 6 and 17 months exposure to altered conditions, with markedly different outcomes. At 6 months, the percentage hatching and larval survival rates were greatest in the animals kept at 0°C under current pH conditions, whilst those under lowered pH and +2°C performed significantly less well. After 17 months, performance was not significantly different across treatments, including controls. However, under the altered conditions urchins produced larger eggs compared with control animals. 5. These data show that under long-term culture adult S. neumayeri appear to acclimate their metabolic and reproductive physiology to the combined stressors of altered pH and increased temperature, with relatively little measureable effect. They also emphasize the importance of long-term studies in evaluating effects of altered pH, particularly in slow developing marine species with long gonad maturation times, as the effects of altered conditions cannot be accurately evaluated unless gonads have fully matured under the new conditions.
According to the size-advantage hypothesis, protandric sequential hermaphroditism is expected when the increase in reproductive success with age or size is small for males but large for females. Interestingly, some protandrous molluscs have developed gregarious strategies that might enhance male reproductive success but at the cost of intraspecific competition. The gastropod Crepidula fornicata, a European invading species, is ideal for investigating mating patterns in a sequential hermaphrodite in relation to grouping behaviour because individuals of different size (age) live in perennial stacks, fertilization is internal and embryos are brooded. Paternity analyses were undertaken in stacks sampled in three close and recently invaded sites in Brittany, France. Paternity assignment of 239 larvae, sampled from a set of 18 brooding females and carried out using five microsatellite loci, revealed that 92% of the crosses occurred between individuals located in the same stack. These stacks thus function as independent mating groups in which individuals may reproduce consecutively as male and female over a short time period, a pattern explained by sperm storage capacity. Gregariousness and sex reversal are promoting reproductive insurance in this species. In addition, females are usually fertilized by several males (78% of the broods were multiply sired) occupying any position within the stack, a result reinforcing the hypothesis of sperm competition. Our study pointed out that mating behaviours and patterns of gender allocation varied in concert across sites suggesting that multiple paternities might enhance sex reversal depending on sperm competition intensity.
International audienceThe reproduction of Crepidula fornicata was studied in the Bay of Brest in order to characterise the first step of the reproductive cycle of this invasive species. The survey was carried out from 2000 to 2003 and different parameters were measured, namely, the percentage of the different sexual stages, the straight length of the shell and the percentage of brooding females using a survey of the embryonic development and the fecundity. The juvenile frequency increases generally from mid-June or mid-August, depending on the year. In 2001 and 2003, a first peak was observed as early as May, but it was followed by a rapid disappearance of the individuals. The sex-ratio female/male increased from 0.22 to 0.46 between 2001 and 2003. The sex change between intermediates and females took place mainly in summer and was well marked in 2001 and 2003. The survey of the embryonic development in the egg capsules brooded by the females provided an annual phenology of the laying and hatching processes. The laying period extends from February to September with three to four major periods of egg-laying per year and corresponding hatching periods about 1 month later. Each female lays two to four times per year on average. The first egg-laying concerned fewer females than subsequent ones, except in 2003, and exhibited a higher fecundity. The annual mean of the number of eggs for each stage was not significantly different, thus indicating no significant mortality rate during embryonic development. For the C. fornicata population in the Bay of Brest, several reproductive characteristics tend to highlight its invasive capacity: (1) a long reproductive period, (2) reproduction in a ‘multi-trials' process equivalent to a spreading out of the risks and (3) a relatively high fecundity
To test the potential of diet switching experiments in ecophysiological studies of marine invertebrates, stable carbon isotope ratios were measured at different seasons in the gonad, adductor muscle, digestive gland and gills of scallops (Pecten maximus) and oysters (Crassostrea gigas) held for 15 days on a constant diet of phytoplankton depleted in 13 C. The aim of this study was to determine if differences in carbon incorporation could be detected among species, seasons and organs, and if so, whether it was consistent with their known energy-allocation patterns. After offering the new diet, isotope values of the different organs gradually shifted and significant differences among organs, seasons and species were found. A carbon incorporation index (CII) was calculated to compare the metabolic activity of each organ of the two species between day 0 and day 15. For both species, the digestive gland had the highest CII, the adductor muscle the lowest, while gonad and gills had intermediate values. The CII was generally much higher in P. maximus than in C. gigas, suggesting higher metabolic activity in this species. Seasonal differences in the CII were also observed for the two species and were interpreted as differences in metabolic activity in accordance with our energy allocation scenario. Therefore, stable isotope diet switching experiments appear to be of great value for assessing metabolic orientation in bivalves.
Defining ecologically relevant upper temperature limits of species is important in the context of environmental change. The approach used in the present paper estimates the relationship between rates of temperature change and upper temperature limits for survival in order to evaluate the maximum long-term survival temperature (Ts). This new approach integrates both the exposure time and the exposure temperature in the evaluation of temperature limits. Using data previously published for different temperate and Antarctic marine environments, we calculated Ts in each environment, which allowed us to calculate a new index: the Warming Allowance (WA). This index is defined as the maximum environmental temperature increase which an ectotherm in a given environment can tolerate, possibly with a decrease in performance but without endangering survival over seasonal or lifetime time-scales. It is calculated as the difference between maximum long-term survival temperature (Ts) and mean maximum habitat temperature. It provides a measure of how close a species, assemblage or fauna are living to their temperature limits for long-term survival and hence their vulnerability to environmental warming. In contrast to data for terrestrial environments showing that warming tolerance increases with latitude, results here for marine environments show a less clear pattern as the smallest WA value was for the Peru upwelling system. The method applied here, relating upper temperature limits to rate of experimental warming, has potential for wide application in the identification of faunas with little capacity to survive environmental warming.
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