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

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.

No evidence of predation causing female-biased sex ratios in marine pelagic copepods

Although sex ratios close to unity are expected in dioecious species, biased sex ratios are common in nature. It is essential to understand causes of skewed sex ratios in situ, as they can lead to mate limitation and have implications for the success of natural populations. Female-skewed sex ratios are commonly observed in copepods in situ. Here we discuss the challenges of copepod sex ratio research and provide a critical review of factors determining copepod sex ratios, focusing on 2 main objectives. The first is a critique of the male predation theory, which is currently the main process thought to be responsible for female-skewed sex ratios. It assumes that males have higher mortality because of increased vulnerability to predation during their search for mates. We show that there is little support for the male predation theory, that sex ratios skewed toward females occur in the absence of predation, that sex ratios are not related to predation pressure, and that where sex-skewed predation does occur, it is biased toward females. Our second objective is to suggest alternative hypotheses regarding the determination of sex ratios. We demonstrate that environmental factors, environmental sex determination and sex change have strong effects on copepod sex ratios, and suggest that differential physiological longevity of males and females may be more important in determining sex ratios than previously thought. We suggest that copepod sex ratios are the result of a mixture of factors.

Female-biased sex ratios in marine pelagic copepods: Response to Hirst et al. (2013)

Hirst et al. (2013; Mar Ecol Prog Ser 489:297-298) suggest that Gusmão et al. (2013; Mar Ecol Prog Ser 482:279-298) misinterpreted the findings of Hirst et al. (2010; Limnol Oceanogr 55:2193-2206). They restate that the major factors determining sex ratio in pelagic copepods act upon the adult stage, but they place less emphasis on the idea that predation on male copepods is a likely determinant, and highlight the role of physiological longevity. Here we reconsider the data and confirm our position that at present there is limited evidence to support the theory of male-skewed predation. However, we agree that sex determination is governed by a combination of factors, with the relative emphasis being the main point of contention between the 2 parties.

Patterns in marine microbial community structure

Programa en Oceanografía

Viruses in the marine environment: community dynamics, phage-host interactions and genomic structure

[EN] There are an estimated 1030 viruses in the world oceans, the majority of which are phages (viruses that infect bacteria). Extensive research has demonstrated the significant influence of marine phages on microbial abundance, community structure, genetic exchange and global biogeochemical cycles. In this thesis, we contribute to increase the knowledge about the ecological role of viruses in marine systems, but also we aimed to provide a better understanding about the interactions between phages and their hosts and the genetic pool and biogeography of some the isolated phages genomes.

Seawater carbonate chemistry, stable carbon isotope fractionation and growth rate during experiments with marine phytoplankton community, 1999

Stable carbon isotope fractionation (%) of 7 marine phytoplankton species grown in different irradiance cycles was measured under nutrient-replete conditions at a high light intensity in batch cultures. Compared to experiments under continuous light, all species exhibited a significantly higher instantaneous growth rate (pi), defined as the rate of carbon fixation during the photo period, when cultivated at 12:12 h. 16:8 h, or 186 h light:dark (L/D) cycles. Isotopic fractionation by the diatoms Skeletonema costatum, Asterionella glacialis, Thalassiosira punctigera, and Coscinodiscus wailesii (Group I) was 4 to 6% lower in a 16:8 h L/D cycle than under continuous light, which we attribute to differences in pi. In contrast, E, in Phaeodactylum tn'cornutum, Thalassiosira weissflogii, and in the dinoflagellate Scrippsiella trochoidea (Group 11) was largely insensitive to day length-related differences in instantaneous growth rate. Since other studies have reported growth-rate dependent fractionation under N-limited conditions in P. tricornutum, pi-related effects on fractionation apparently depend on the factor controlling growth rate. We suggest that a general relationship between E, and pi/[C02,,,] may not exist. For 1 species of each group we tested the effect of variable CO2 concentration, [COz,,,], on isotopic fractionation. A decrease in [CO2,,,] from ca 26 to 3 pm01 kg-' caused a decrease in E, by less than 3%0 This indicates that variation in h in response to changes in day length has a similar or even greater effect on isotopic fractionation than [COz,,,] m some of the species tested. In both groups E, tended to be higher in smaller species at comparable growth rates. In 24 and 48 h time series the algal cells became progressively enriched in 13C during the day and the first hours of the dark period, followed by l3C depletion in the 2 h before beginning of the following Light period. The daily amplitude of the algal isotopic composition (613C), however, was <1.5%0, which demonstrates that diurnal variation in Fl3C is relatively small.

Data compilation of respiration, feeding, and growth rates of marine pelagic organisms

The metabolic rate of organisms may either be viewed as a basic property from which other vital rates and many ecological patterns emerge and that follows a universal allometric mass scaling law; or it may be considered a property of the organism that emerges as a result of the organism's adaptation to the environment, with consequently less universal mass scaling properties. Data on body mass, maximum ingestion and clearance rates, respiration rates and maximum growth rates of animals living in the ocean epipelagic were compiled from the literature, mainly from original papers but also from previous compilations by other authors. Data were read from tables or digitized from graphs. Only measurements made on individuals of know size, or groups of individuals of similar and known size were included. We show that clearance and respiration rates have life-form-dependent allometries that have similar scaling but different elevations, such that the mass-specific rates converge on a rather narrow size-independent range. In contrast, ingestion and growth rates follow a near-universal taxa-independent ~3/4 mass scaling power law. We argue that the declining mass-specific clearance rates with size within taxa is related to the inherent decrease in feeding efficiency of any particular feeding mode. The transitions between feeding mode and simultaneous transitions in clearance and respiration rates may then represent adaptations to the food environment and be the result of the optimization of tradeoffs that allow sufficient feeding and growth rates to balance mortality.

Small Changes in pH Have Direct Effects on Marine Bacterial Community Composition: A Microcosm Approach

As the atmospheric CO2 concentration rises, more CO2 will dissolve in the oceans, leading to a reduction in pH. Effects of ocean acidification on bacterial communities have mainly been studied in biologically complex systems, in which indirect effects, mediated through food web interactions, come into play. These approaches come close to nature but suffer from low replication and neglect seasonality. To comprehensively investigate direct pH effects, we conducted highly-replicated laboratory acidification experiments with the natural bacterial community from Helgoland Roads (North Sea). Seasonal variability was accounted for by repeating the experiment four times (spring, summer, autumn, winter). Three dilution approaches were used to select for different ecological strategies, i.e. fast-growing or low-nutrient adapted bacteria. The pH levels investigated were in situ seawater pH (8.15-8.22), pH 7.82 and pH 7.67, representing the present-day situation and two acidification scenarios projected for the North Sea for the year 2100. In all seasons, both automated ribosomal intergenic spacer analysis and 16S ribosomal amplicon pyrosequencing revealed pH-dependent community shifts for two of the dilution approaches. Bacteria susceptible to changes in pH were different members of Gammaproteobacteria, Flavobacteriaceae, Rhodobacteraceae, Campylobacteraceae and further less abundant groups. Their specific response to reduced pH was often context-dependent. Bacterial abundance was not influenced by pH. Our findings suggest that already moderate changes in pH have the potential to cause compositional shifts, depending on the community assembly and environmental factors. By identifying pH-susceptible groups, this study provides insights for more directed, in-depth community analyses in large-scale and long-term experiments.

Short- and long-term conditioning of a temperate marine diatom community to acidification and warming

Ocean acidification and greenhouse warming will interactively influence competitive success of key phytoplankton groups such as diatoms, but how long-term responses to global change will affect community structure is unknown. We incubated a mixed natural diatom community from coastal New Zealand waters in a short-term (two-week) incubation experiment using a factorial matrix of warming and/or elevated pCO2 and measured effects on community structure. We then isolated the dominant diatoms in clonal cultures and conditioned them for 1 year under the same temperature and pCO2 conditions from which they were isolated, in order to allow for extended selection or acclimation by these abiotic environmental change factors in the absence of interspecific interactions. These conditioned isolates were then recombined into 'artificial' communities modelled after the original natural assemblage and allowed to compete under conditions identical to those in the short-term natural community experiment. In general, the resulting structure of both the unconditioned natural community and conditioned 'artificial' community experiments was similar, despite differences such as the loss of two species in the latter. pCO2 and temperature had both individual and interactive effects on community structure, but temperature was more influential, as warming significantly reduced species richness. In this case, our short-term manipulative experiment with a mixed natural assemblage spanning weeks served as a reasonable proxy to predict the effects of global change forcing on diatom community structure after the component species were conditioned in isolation over an extended timescale. Future studies will be required to assess whether or not this is also the case for other types of algal communities from other marine regimes.

Data compilation of marine pelagic organism biomasses, swimming velocities, clearance and respiration rates

Jellyfishes have functionally replaced several overexploited commercial stocks of planktivorous fishes. This is paradoxical, because they use a primitive prey capture mechanism requiring direct contact with the prey, whereas fishes use more efficient visual detection. We have compiled published data to show that, in spite of their primitive life-style, jellyfishes exhibit similar instantaneous prey clearance and respiration rates as their fish competitors and similar potential for growth and reproduction. To achieve this production, they have evolved large, water-laden bodies that increase prey contact rates. Although larger bodies are less efficient for swimming, optimization analysis reveals that large collectors are advantageous if they move through the water sufficiently slowly.