Female-biased sex ratios in marine pelagic copepods: comment on Gusmao et al

Gusmão et al. (2013; Mar Ecol Prog Ser 482:279-298) review causes of sex ratio skew in pelagic copepods and in doing so repeatedly dispute the paper of Hirst et al. (2010) ‘Does predation control adult sex ratios and longevities in marine pelagic copepods?’ Here we respond to some important errors in their citation of our paper and briefly highlight where future work is needed in order to attribute the causes of strong sex ratio skew seen in some copepod families.

Reproductive effects of endocrine disrupting chemicals, bisphenol-A and 17β-oestradiol, on Cerastoderma edule from south-west England: field study and laboratory exposure

Endocrine disruption has rarely been reported in field populations of the edible cockle and the context with the general health of the shellfish is unclear. This study examined the reproductive state of two Cerastoderma edule populations over a 6-month period to assess their reproductive condition, the incidence of intersex and presence of parasitic infection. A further seven native sites from south-west England were examined during the peak reproductive season to identify the presence of intersex within the region. Laboratory exposures of organisms collected from field populations showed a significantly female-biased sex ratio compared with controls when exposed to the endocrine disrupting chemicals, bisphenol-A (nominal concentration: 0.1 µg L−1) and 17β-oestradiol (nominal concentration: 0.1 µg L−1), but none of the chemical exposures induced intersex. Intersex was revealed in seven out of the nine native populations of C. edule sampled at peak reproductive season. The highest incidence and most severe case of intersex were reported at Lower Anderton on the River Tamer which also had a significantly female-biased sex ratio. Additionally, the dominant trematode family was the Bucephalaidae. Parasitic infection influences the maturity of C. edule by lowering both mean gonad index and condition index. These results suggest that endocrine disrupting chemicals could be contributing factors towards the development of intersex in C. edule.

Reproductive effects of endocrine disrupting chemicals, bisphenol-A and 17β-oestradiol, on Cerastoderma edule from south-west England: field study and laboratory exposure

Endocrine disruption has rarely been reported in field populations of the edible cockle and the context with the general health of the shellfish is unclear. This study examined the reproductive state of two Cerastoderma edule populations over a 6-month period to assess their reproductive condition, the incidence of intersex and presence of parasitic infection. A further seven native sites from south-west England were examined during the peak reproductive season to identify the presence of intersex within the region. Laboratory exposures of organisms collected from field populations showed a significantly female-biased sex ratio compared with controls when exposed to the endocrine disrupting chemicals, bisphenol-A (nominal concentration: 0.1 µg L−1) and 17β-oestradiol (nominal concentration: 0.1 µg L−1), but none of the chemical exposures induced intersex. Intersex was revealed in seven out of the nine native populations of C. edule sampled at peak reproductive season. The highest incidence and most severe case of intersex were reported at Lower Anderton on the River Tamer which also had a significantly female-biased sex ratio. Additionally, the dominant trematode family was the Bucephalaidae. Parasitic infection influences the maturity of C. edule by lowering both mean gonad index and condition index. These results suggest that endocrine disrupting chemicals could be contributing factors towards the development of intersex in C. edule.

Does predation control adult sex ratios and longevities in marine pelagic copepods?

We assess the causes of adult sex ratio skew in marine pelagic copepods by examining changes in these ratios between the juveniles and adults, sexual differences in juvenile stage durations, and mortality rates of adults in the field and laboratory (when free from predators). In the field, late copepodite stages (CIV and CV) commonly have sex ratios that are either not significantly different from equity (1 : 1), or slightly male biased. By contrast, in adults, these ratios are commonly significantly biased toward female dominance. Sex ratio skews are therefore primarily attributable to processes in adults. Members of the non-Diaptomoidea have especially skewed adult ratios; in the members Oithonidae and Clausocalanidae this is not generated from differences between male and female adult physiological longevity (i.e., laboratory longevity when free of predators). In the genera Acartia, Oithona, and Pseudocalanus, we estimate that predation mortality contributed ≥ 69% of the field mortality rate in adult males, whereas in Acartia, Oithona, and Calanus adult females, this is ≥ 36%.We conclude that (1) adult sex ratio skew in pelagic copepods is primarily due to differential mortality of the sexes in the adult stage and not in juveniles, (2) mortality rates of adult Acartia, Pseudocalanus, and Oithona are dominated by predation mortality rather than physiological longevity (except under extreme food limitation), and (3) in Pseudocalanus and Oithona, elevated mortality rates in adult males to females is predominantly due to higher predation on males. Our work demonstrates that we now need to develop a more comprehensive understanding of the importance of feeding preferences in predators. Continue reading full article

Broad-scale patterns of sex ratios in Patella spp.: a comparison of range edge and central range populations in the British Isles and Portugal

Sex change, or sequential hermaphroditism, occurs in the plant and animal kingdoms and often determines a predominance of the first sex. Our aim was to explore changes in sex ratios within the range of the species studied: Patella vulgata and Patella depressa. The broad-scale survey of sex with size of limpets covered a range of latitudes from Zambujeira do Mar (southern Portugal) to the English Channel. Indirect evidence was found for the occurrence of protandry in P. vulgata populations from the south of England, with females predominating in larger size-classes; cumulative frequency distributions of males and females were different; sex ratios were biased towards males and smallest sizes of males were smaller than the smallest sizes of females. In contrast in Portugal females were found in most size-classes of P. vulgata. In P. depressa populations from the south coast of England and Portugal females were interspersed across most size-classes; size distributions of males and females and size at first maturity of males and females did not differ. P. depressa did, however, show some indications of the possibility of slight protandry occurring in Portugal. The test of sex ratio variation with latitude indicated that P. vulgata sex ratios might be involved in determining the species range limit, particularly at the equatorward limit since the likelihood of being male decreased from the south coast of England to southern Portugal. Thus at the southern range limit, sperm could be in short supply due to scarcity of males contributing to an Allee effect.

Broad-scale patterns of sex ratios in Patella spp.: a comparison of range edge and central range populations in the British Isles and Portugal

Sex change, or sequential hermaphroditism, occurs in the plant and animal kingdoms and often determines a predominance of the first sex. Our aim was to explore changes in sex ratios within the range of the species studied: Patella vulgata and Patella depressa. The broad-scale survey of sex with size of limpets covered a range of latitudes from Zambujeira do Mar (southern Portugal) to the English Channel. Indirect evidence was found for the occurrence of protandry in P. vulgata populations from the south of England, with females predominating in larger size-classes; cumulative frequency distributions of males and females were different; sex ratios were biased towards males and smallest sizes of males were smaller than the smallest sizes of females. In contrast in Portugal females were found in most size-classes of P. vulgata. In P. depressa populations from the south coast of England and Portugal females were interspersed across most size-classes; size distributions of males and females and size at first maturity of males and females did not differ. P. depressa did, however, show some indications of the possibility of slight protandry occurring in Portugal. The test of sex ratio variation with latitude indicated that P. vulgata sex ratios might be involved in determining the species range limit, particularly at the equatorward limit since the likelihood of being male decreased from the south coast of England to southern Portugal. Thus at the southern range limit, sperm could be in short supply due to scarcity of males contributing to an Allee effect.

The influence of simulated exploitation onPatella vulgatapopulations: protandric sex change is size-dependent

Grazing mollusks are used as a food resource worldwide, and limpets are harvested commercially for both local consumption and export in several countries. This study describes a field experiment to assess the effects of simulated human exploitation of limpets Patella vulgata on their population ecology in terms of protandry (age-related sex change from male to female), growth, recruitment, migration, and density regulation. Limpet populations at two locations in southwest England were artificially exploited by systematic removal of the largest individuals for 18 months in plots assigned to three treatments at each site: no (control), low, and high exploitation. The shell size at sex change (L50: the size at which there is a 50:50 sex ratio) decreased in response to the exploitation treatments, as did the mean shell size of sexual stages. Size-dependent sex change was indicated by L50 occurring at smaller sizes in treatments than controls, suggesting an earlier switch to females. Mean shell size of P. vulgata neuters changed little under different levels of exploitation, while males and females both decreased markedly in size with exploitation. No differences were detected in the relative abundances of sexual stages, indicating some compensation for the removal of the bigger individuals via recruitment and sex change as no migratory patterns were detected between treatments. At the end of the experiment, 0–15 mm recruits were more abundant at one of the locations but no differences were detected between treatments. We conclude that sex change in P. vulgata can be induced at smaller sizes by reductions in density of the largest individuals reducing interage class competition. Knowledge of sex-change adaptation in exploited limpet populations should underpin strategies to counteract population decline and improve rocky shore conservation and resource management.

The influence of simulated exploitation onPatella vulgatapopulations: protandric sex change is size-dependent

Grazing mollusks are used as a food resource worldwide, and limpets are harvested commercially for both local consumption and export in several countries. This study describes a field experiment to assess the effects of simulated human exploitation of limpets Patella vulgata on their population ecology in terms of protandry (age-related sex change from male to female), growth, recruitment, migration, and density regulation. Limpet populations at two locations in southwest England were artificially exploited by systematic removal of the largest individuals for 18 months in plots assigned to three treatments at each site: no (control), low, and high exploitation. The shell size at sex change (L50: the size at which there is a 50:50 sex ratio) decreased in response to the exploitation treatments, as did the mean shell size of sexual stages. Size-dependent sex change was indicated by L50 occurring at smaller sizes in treatments than controls, suggesting an earlier switch to females. Mean shell size of P. vulgata neuters changed little under different levels of exploitation, while males and females both decreased markedly in size with exploitation. No differences were detected in the relative abundances of sexual stages, indicating some compensation for the removal of the bigger individuals via recruitment and sex change as no migratory patterns were detected between treatments. At the end of the experiment, 0–15 mm recruits were more abundant at one of the locations but no differences were detected between treatments. We conclude that sex change in P. vulgata can be induced at smaller sizes by reductions in density of the largest individuals reducing interage class competition. Knowledge of sex-change adaptation in exploited limpet populations should underpin strategies to counteract population decline and improve rocky shore conservation and resource management.

Growth and shrinkage in Antarctic krill Euphausia superba is sex-dependent

ABSTRACT: The ability of Antarctic krill Euphausia superba Dana to withstand the overwintering period is critical to their success. Laboratory evidence suggests that krill may shrink in body length during this time in response to the low availability of food. Nevertheless, verification that krill can shrink in the natural environment is lacking because winter data are difficult to obtain. One of the few sources of winter krill population data is from commercial vessels. We examined length-frequency data of adult krill (>35 mm total body length) obtained from commercial vessels in the Scotia-Weddell region and compared our results with those obtained from a combination of science and commercial sampling operations carried out in this region at other times of the year. Our analyses revealed body-length shrinkage in adult females but not males during overwinter, based on both the tracking of modal size classes over seasons and sex-ratio patterns. Other explanatory factors, such as differential mortality, immigration and emigration, could not explain the observed differences. The same pattern was also observed at South Georgia and in the Western Antarctic Peninsula. Fitted seasonally modulated von Bertalanffy growth functions predicted a pattern of overwintering shrinkage in all body-length classes of females, but only stagnation in growth in males. This shrinkage most likely reflects morphometric changes resulting from the contraction of the ovaries and is not necessarily an outcome of winter hardship. The sex-dependent changes that we observed need to be incorporated into life cycle and population dynamic models of this species, particularly those used in managing the fishery. KEY WORDS: Southern Ocean · Population dynamics · Production · Life cycle · Fishery

Growth and shrinkage in Antarctic krill Euphausia superba is sex-dependent

ABSTRACT: The ability of Antarctic krill Euphausia superba Dana to withstand the overwintering period is critical to their success. Laboratory evidence suggests that krill may shrink in body length during this time in response to the low availability of food. Nevertheless, verification that krill can shrink in the natural environment is lacking because winter data are difficult to obtain. One of the few sources of winter krill population data is from commercial vessels. We examined length-frequency data of adult krill (>35 mm total body length) obtained from commercial vessels in the Scotia-Weddell region and compared our results with those obtained from a combination of science and commercial sampling operations carried out in this region at other times of the year. Our analyses revealed body-length shrinkage in adult females but not males during overwinter, based on both the tracking of modal size classes over seasons and sex-ratio patterns. Other explanatory factors, such as differential mortality, immigration and emigration, could not explain the observed differences. The same pattern was also observed at South Georgia and in the Western Antarctic Peninsula. Fitted seasonally modulated von Bertalanffy growth functions predicted a pattern of overwintering shrinkage in all body-length classes of females, but only stagnation in growth in males. This shrinkage most likely reflects morphometric changes resulting from the contraction of the ovaries and is not necessarily an outcome of winter hardship. The sex-dependent changes that we observed need to be incorporated into life cycle and population dynamic models of this species, particularly those used in managing the fishery. KEY WORDS: Southern Ocean · Population dynamics · Production · Life cycle · Fishery

The Influence Of Temperature And Salinity On Energy Partitioning In The Marine Nematode Diplolaimelloides-Bruciei

Measurements of population growth, generation time, fecundity and respiration in laboratory culture have been made, in relation to temperature and salinity, for the nematode Diplolaimelloides bruciei Hopper, a species normally associated with decayed material of the marsh grass Spartina. The intrinsic rate of increase (r) is high: it is related to temperature between 5° and 25°C by a sigmoid function which is steepest between 10° and 15°C, and is maximum at 26‰ salinity. Generation time is related to temperature by a power function and is shortest at 26‰ salinity. The effect of temperature on generation time is consistent with other data for marine nematodes, and the steep slope of r against temperature is largely due to the marked effect of temperature on fecundity. A sex ratio of 2:1 in favour of males is maintained regardless of culture conditions or population density. Respiration increases exponentially with temperature between 5° and 25°C, with a very high Q10 (3.94), but is not affected by salinity. At 30°C respiration is no higher than at 25°C. A high and relatively stable production efficiency (P/A) is maintained between 10 and 30°C with a maximum of 87% at 15°C; there is a stable reproductive effort (Pr/A) of about 10%. At 5°C both these ratios are zero. Data for the harpacticoid copepod Tachidius discipes, derived from the literature, show that this too has a high and stable production efficiency, which may be a characteristic of meiofaunal species in general, but in this species efficiency is relatively high at 5°C. Many features of the energy balance in D. bruciei can be related to an opportunistic mode of life.