Lunar rhythms in growth of larval fish.

Growth and survival of larval fishes is highly variable and unpredictable. Our limited understanding of this variation constrains our ability to forecast population dynamics and effectively manage fisheries. Here we show that daily growth rates of a coral reef fish (the sixbar wrasse, Thalassoma hardwicke) are strongly lunar-periodic and predicted by the timing of nocturnal brightness: growth was maximized when the first half of the night was dark and the second half of the night was bright. Cloud cover that obscured moonlight facilitated a 'natural experiment', and confirmed the effect of moonlight on growth. We suggest that lunar-periodic growth may be attributable to light-mediated suppression of diel vertical migrations of predators and prey. Accounting for such effects will improve our capacity to predict the future dynamics of marine populations, especially in response to climate-driven changes in nocturnal cloud cover and intensification of artificial light, which could lead to population declines by reducing larval survival and growth.

Spatial aggregation of aquatic habitats affects oviposition patterns in Aedes mosquitoes.

Colonization, including oviposition, is an important driver of population and community dynamics both within and across habitat patches. Most research has focused on the roles of habitat availability or quality on colonization and its outcomes. However, the spatial distribution of habitats also likely affects these processes. We conducted field experiments in Georgia, USA, using clustered and dispersed arrays of equal numbers of oviposition patches to investigate how patch aggregation influenced oviposition by Aedes mosquitoes. We tested the effects of aggregation on: (1) the total number of eggs an array received, (2) the proportion of patches within an array that received eggs, and (3) the number of eggs per colonized patch. We compared results to predictions from three models (Field of Dreams, Propagule Redirection, and Excess Attraction), which vary in the degree to which arrays attract colonists and apportion those colonists among patches. Clustered arrays received 22% more eggs than dispersed arrays, with clustered patches significantly more likely to receive eggs. At the species level, A. albopictus responded more to clustering than did A. triseriatus. These results are inconsistent with Propagule Redirection, but support the Excess Attraction and Field of Dreams models. Although clustered arrays occupied a relatively small area, they attracted at least as many ovipositing mosquitoes as did dispersed arrays. However, the number of eggs per colonized patch did not differ between clustered and dispersed arrays. Therefore, density dependence among larvae, and hence the production of adult mosquitoes on a per-patch basis, should be similar in dispersed and clustered landscapes.

Habitat-dependent movement rate can determine the efficacy of marine protected areas.

Theoretical studies of marine protected areas (MPAs) suggest that more mobile species should exhibit reduced local effects (defined as the ratio of the density inside vs. outside of the MPA). However, empirical studies have not supported the expected negative relationship between the local effect and mobility. We propose that differential, habitat-dependent movement (i.e., a higher movement rate in the fishing grounds than in the MPA) might explain the disparity between theoretical expectations and empirical results. We evaluate this hypothesis by building two-patch box and stepping-stone models and show that increasing disparity in the habitat-specific movement rates shifts the relationship between the local effect and mobility from negative (the previous theoretical results) to neutral or positive (the empirical pattern). This shift from negative to positive occurs when differential movement offsets recruitment and mortality differences between the two habitats. Thus, local effects of MPAs might be caused by behavioral responses via differential movement rather than by, or in addition to, reductions in mortality. In addition, the benefits of MPAs, in terms of regional abundance and fishing yields, can be altered by the magnitude of differential movement. Thus, our study points to a need for empirical investigations that disentangle the interactions among mobility, differential movement, and protection.

Faster turnover of new soil carbon inputs under increased atmospheric CO2.

Rising levels of atmospheric CO2 frequently stimulate plant inputs to soil, but the consequences of these changes for soil carbon (C) dynamics are poorly understood. Plant-derived inputs can accumulate in the soil and become part of the soil C pool ("new soil C"), or accelerate losses of pre-existing ("old") soil C. The dynamics of the new and old pools will likely differ and alter the long-term fate of soil C, but these separate pools, which can be distinguished through isotopic labeling, have not been considered in past syntheses. Using meta-analysis, we found that while elevated CO2 (ranging from 550 to 800 parts per million by volume) stimulates the accumulation of new soil C in the short term (<1 year), these effects do not persist in the longer term (1-4 years). Elevated CO2 does not affect the decomposition or the size of the old soil C pool over either temporal scale. Our results are inconsistent with predictions of conventional soil C models and suggest that elevated CO2 might increase turnover rates of new soil C. Because increased turnover rates of new soil C limit the potential for additional soil C sequestration, the capacity of land ecosystems to slow the rise in atmospheric CO2 concentrations may be smaller than previously assumed.

When environmental factors become stressors: interactive effects of vermetid gastropods and sedimentation on corals.

Environmental stressors often interact, but most studies of multiple stressors have focused on combinations of abiotic stressors. Here we examined the potential interaction between a biotic stressor, the vermetid snail Ceraesignum maximum, and an abiotic stressor, high sedimentation, on the growth of reef-building corals. In a field experiment, we subjected juvenile massive Porites corals to four treatments: (i) neither stressor, (ii) sedimentation, (iii) vermetids or (iv) both stressors. Unexpectedly, we found no effect of either stressor in isolation, but a significant decrease in coral growth in the presence of both stressors. Additionally, seven times more sediment remained on corals in the presence (versus absence) of vermetids, likely owing to adhesion of sediments to corals via vermetid mucus. Thus, vermetid snails and high sedimentation can interact to drive deleterious effects on reef-building corals. More generally, our study illustrates that environmental factors can combine to have negative interactive effects even when individual effects are not detectable. Such 'ecological surprises' may be easily overlooked, leading to environmental degradation that cannot be anticipated through the study of isolated factors.

Increased soil emissions of potent greenhouse gases under increased atmospheric CO2.

Increasing concentrations of atmospheric carbon dioxide (CO(2)) can affect biotic and abiotic conditions in soil, such as microbial activity and water content. In turn, these changes might be expected to alter the production and consumption of the important greenhouse gases nitrous oxide (N(2)O) and methane (CH(4)) (refs 2, 3). However, studies on fluxes of N(2)O and CH(4) from soil under increased atmospheric CO(2) have not been quantitatively synthesized. Here we show, using meta-analysis, that increased CO(2) (ranging from 463 to 780 parts per million by volume) stimulates both N(2)O emissions from upland soils and CH(4) emissions from rice paddies and natural wetlands. Because enhanced greenhouse-gas emissions add to the radiative forcing of terrestrial ecosystems, these emissions are expected to negate at least 16.6 per cent of the climate change mitigation potential previously predicted from an increase in the terrestrial carbon sink under increased atmospheric CO(2) concentrations. Our results therefore suggest that the capacity of land ecosystems to slow climate warming has been overestimated.

Enrichment scale determines herbivore control of primary producers.

Anthropogenic nutrient enrichment stimulates primary production and threatens natural communities worldwide. Herbivores may counteract deleterious effects of enrichment by increasing their consumption of primary producers. However, field tests of herbivore control are often done by adding nutrients at small (e.g., sub-meter) scales, while enrichment in real systems often occurs at much larger scales (e.g., kilometers). Therefore, experimental results may be driven by processes that are not relevant at larger scales. Using a mathematical model, we show that herbivores can control primary producer biomass in experiments by concentrating their foraging in small enriched plots; however, at larger, realistic scales, the same mechanism may not lead to herbivore control of primary producers. Instead, other demographic mechanisms are required, but these are not examined in most field studies (and may not operate in many systems). This mismatch between experiments and natural processes suggests that many ecosystems may be less resilient to degradation via enrichment than previously believed.

Application of a two-pool model to soil carbon dynamics under elevated CO2.

Elevated atmospheric CO2 concentrations increase plant productivity and affect soil microbial communities, with possible consequences for the turnover rate of soil carbon (C) pools and feedbacks to the atmosphere. In a previous analysis (Van Groenigen et al., 2014), we used experimental data to inform a one-pool model and showed that elevated CO2 increases the decomposition rate of soil organic C, negating the storage potential of soil. However, a two-pool soil model can potentially explain patterns of soil C dynamics without invoking effects of CO2 on decomposition rates. To address this issue, we refit our data to a two-pool soil C model. We found that CO2 enrichment increases decomposition rates of both fast and slow C pools. In addition, elevated CO2 decreased the carbon use efficiency of soil microbes (CUE), thereby further reducing soil C storage. These findings are consistent with numerous empirical studies and corroborate the results from our previous analysis. To facilitate understanding of C dynamics, we suggest that empirical and theoretical studies incorporate multiple soil C pools with potentially variable decomposition rates.

Optimal sampling strategies for detecting zoonotic disease epidemics.

The early detection of disease epidemics reduces the chance of successful introductions into new locales, minimizes the number of infections, and reduces the financial impact. We develop a framework to determine the optimal sampling strategy for disease detection in zoonotic host-vector epidemiological systems when a disease goes from below detectable levels to an epidemic. We find that if the time of disease introduction is known then the optimal sampling strategy can switch abruptly between sampling only from the vector population to sampling only from the host population. We also construct time-independent optimal sampling strategies when conducting periodic sampling that can involve sampling both the host and the vector populations simultaneously. Both time-dependent and -independent solutions can be useful for sampling design, depending on whether the time of introduction of the disease is known or not. We illustrate the approach with West Nile virus, a globally-spreading zoonotic arbovirus. Though our analytical results are based on a linearization of the dynamical systems, the sampling rules appear robust over a wide range of parameter space when compared to nonlinear simulation models. Our results suggest some simple rules that can be used by practitioners when developing surveillance programs. These rules require knowledge of transition rates between epidemiological compartments, which population was initially infected, and of the cost per sample for serological tests.

Coral guard crabs.

Adrian Stier and Craig Osenberg introduce Trapeziid crabs, which live in close symbiosis with corals.

Mosquito population dynamics is shaped by the interaction among larval density, season, and land use.

Understanding how variation in key abiotic and biotic factors interact at spatial scales relevant for mosquito fitness and population dynamics is crucial for predicting current and future mosquito distributions and abundances, and the transmission potential for human pathogens. However, studies investigating the effects of environmental variation on mosquito traits have investigated environmental factors in isolation or in laboratory experiments that examine constant environmental conditions that often do not occur in the field. To address these limitations, we conducted a semi-field experiment in Athens, Georgia using the invasive Asian tiger mosquito (Aedes albopictus). We selected nine sites that spanned natural variation in impervious surface and vegetation cover to explore effects of the microclimate (temperature and humidity) on mosquitoes. On these sites, we manipulated conspecific larval density at each site. We repeated the experiment in the summer and fall. We then evaluated the effects of land cover, larval density, and time of season, as well as interactive effects, on the mean proportion of females emerging, juvenile development time, size upon emergence, and predicted per capita population growth (i.e., fitness). We found significant effects of larval density, land cover, and season on all response variables. Of most note, we saw strong interactive effects of season and intra-specific density on each response variable, including a non-intuitive decrease in development time with increasing intra-specific competition in the fall. Our study demonstrates that ignoring the interaction between variation in biotic and abiotic variables could reduce the accuracy and precision of models used to predict mosquito population and pathogen transmission dynamics, especially those inferring dynamics at finer-spatial scales across which transmission and control occur.

Para poder predecir la distribución y abundancia de las poblaciones de mosquitos y la transmisión potencial de patógenos a humanos, es crucial comprender cómo factores abióticos y bióticos clave para el éxito reproductivo y la dinámica poblacional de los mosquitos interactúan a escalas relevantes. Sin embargo, los estudios que han investigado los efectos de variables ambientales en las características demográficas de los mosquitos han considerado su efecto de forma aislada o en experimentos de laboratorio bajo condiciones ambientales constantes que, a menudo, no reflejan lo que ocurre en el campo. Para abordar estas limitaciones, llevamos a cabo un experimento de semi-campo en Athens, Georgia, utilizando el mosquito invasor tigre asiático (Aedes albopictus). Seleccionamos nueve sitios que abarcaban variaciones naturales en la superficie impermeable y cobertura vegetal para explorar los efectos del microclima (temperatura y humedad) en los mosquitos. También manipulamos la densidad de larvas de tigre asiático en dos experimentos que fueron realizados en el verano y otoño. Evaluamos los efectos de la cobertura vegetal, la densidad de larvas, la temporada climática, y la interacción entre estas variables en la proporción de hembras que emergieron, el tiempo de desarrollo de las larvas, el tamaño al momento de la emergencia, y el crecimiento demográfico per cápita previsto (éxito reproductivo). Encontramos efectos significativos de la densidad de larvas, la variación de la cobertura vegetal y la estación del año en todas las variables de respuesta. Más notablemente, observamos un fuerte efecto de la interacción entre la temporada climática y la densidad de larvas en todas las variables de respuesta, incluyendo una disminución no intuitiva en el tiempo de desarrollo con el aumento de la competencia intraespecífica en el otoño. Nuestro estudio evidencia que ignorar la interacción entre variables abióticas y bióticas podría reducir la exactitud y precisión de los modelos utilizados para predecir las dinámicas de las poblaciones de mosquitos, y por tanto, de la transmisión de patógenos. Esto, especialmente en modelos que infieren estas dinámicas a escalas espaciales más finas, en las cuales ocurre la transmisión y el control.

A meta-evaluation of the quality of reporting and execution in ecological meta-analyses.

Quantitatively summarizing results from a collection of primary studies with meta-analysis can help answer ecological questions and identify knowledge gaps. The accuracy of the answers depends on the quality of the meta-analysis. We reviewed the literature assessing the quality of ecological meta-analyses to evaluate current practices and highlight areas that need improvement. From each of the 18 review papers that evaluated the quality of meta-analyses, we calculated the percentage of meta-analyses that met criteria related to specific steps taken in the meta-analysis process (i.e., execution) and the clarity with which those steps were articulated (i.e., reporting). We also re-evaluated all the meta-analyses available from Pappalardo et al. [1] to extract new information on ten additional criteria and to assess how the meta-analyses recognized and addressed non-independence. In general, we observed better performance for criteria related to reporting than for criteria related to execution; however, there was a wide variation among criteria and meta-analyses. Meta-analyses had low compliance with regard to correcting for phylogenetic non-independence, exploring temporal trends in effect sizes, and conducting a multifactorial analysis of moderators (i.e., explanatory variables). In addition, although most meta-analyses included multiple effect sizes per study, only 66% acknowledged some type of non-independence. The types of non-independence reported were most often related to the design of the original experiment (e.g., the use of a shared control) than to other sources (e.g., phylogeny). We suggest that providing specific training and encouraging authors to follow the PRISMA EcoEvo checklist recently developed by O'Dea et al. [2] can improve the quality of ecological meta-analyses.

Emergent effects of multiple predators on prey survival: the importance of depletion and the functional response.

The combined effects of multiple predators often cannot be predicted from their independent effects. Emergent multiple predator effects (MPEs) include risk enhancement, where combined predators kill more prey than predicted by their individual effects, and risk reduction, where fewer prey are killed than predicted. Current methods for detecting MPEs are biased because they assume linear functional responses and/or no prey depletion. As a result, past studies overestimated the occurrence of risk enhancement for additive designs, and tended to overestimate the occurrence of risk reduction for substitutive designs. Characterising the predators' functional responses and accounting for prey depletion reduces biases in detection, estimation, interpretation and generalisation of the emergent effects of predator diversity on prey survival. These findings have implications beyond MPE's and should be considered in all studies aimed at understanding how multiple factors combine when demographic rates are density dependent.

Differential movement and movement bias models for marine protected areas.

Marine protected areas (MPAs) are promoted as a tool to protect overfished stocks and increase fishery yields. Previous models suggested that adult mobility modified effects of MPAs by reducing densities of fish inside reserves, but increasing yields (i.e., increasing densities outside of MPAs). Empirical studies contradicted this prediction: as mobility increased, the relative density of fishes inside MPAs (relative to outside) increased or stayed constant. We hypothesized that this disparity between theoretical and empirical results was the result of differential movement of fish inside versus outside the MPA. We, therefore, developed a model with unequal and discontinuous diffusion, and analyzed its steady state and stability. We determined the abundance in the fishing grounds, the yield, the total abundance and the log ratio at steady-state and examined their response to adult mobility (while keeping the relative inequity in the diffusion constant). Abundance in the fishing grounds and yield increased, while total abundance and log-ratio decreased, as mobility increased. These results were all qualitatively consistent with the previous models assuming uniform diffusivity. Thus, the mismatch between empirical and theoretical results must result from other processes or other forms of differential movement. Therefore, we modified our original model by assuming that species located on the boundary of the MPA will preferentially move towards the MPA. This localized movement bias model gives rise to steady state profiles that can differ radically from the profiles in the unbiased model, especially when the bias is large. Moreover, for sufficiently large bias values, the monotonicity of the four measures with increased mobility is reversed, when compared with our original model. Thus, the movement bias model reconciles empirical data and theoretical results.

Thermal Traits Vary with Mass and across Populations of the Marsh Periwinkle, Littoraria irrorata.

AbstractPhysiological processes influence how individuals perform in various environmental contexts. The basis of such processes, metabolism, scales allometrically with body mass and nonlinearly with temperature, as described by a thermal performance curve. Past studies of thermal performance curves tend to focus on effects of temperature on a single body size or population, rather than variation in the thermal performance curve across sizes and populations. Here, we estimate intraspecific variation in parameters of the thermal performance curve in the salt marsh gastropod Littoraria irrorata. First, we quantify the thermal performance curve for respiration rate as a function of both temperature and body size in Littoraria and evaluate whether the thermal parameters and body size scaling are interdependent. Next, we quantify how parameters in the thermal performance curve for feeding rate vary between three Littoraria populations that occur along a latitudinal gradient. Our work suggests that the thermal traits describing Littoraria respiration are dependent on body mass and that both the thermal traits and the mass scaling of feeding vary across sites. We found limited evidence to suggest that mass scaling of Littoraria feeding or respiration rates depends on temperature. Variation in the thermal performance curves interacts with the size structure of the Littoraria population to generate divergent population-level responses to temperature. These results highlight the importance of considering variation in population size structure and physiological allometry when attempting to predict how temperature change will affect physiological responses and consumer-resource interactions.

How moonlight shapes environments, life histories, and ecological interactions on coral reefs.

The lunar cycle drives variation in nocturnal brightness. For the epipelagic larvae of coral reef organisms, nocturnal illumination may have widespread and underappreciated consequences. At sea, the onset of darkness coincides with an influx of mesopelagic organisms to shallow water (i.e. 'diel vertical migrants') that include predators (e.g. lanternfishes) and prey (zooplankton) of zooplanktivorous coral reef larvae. Moonlight generally suppresses this influx, but lunar periodicity in the timing and intensity of nocturnal brightness may affect vertically migrating predators and prey differently. A major turnover of species occurs at sunset on the reef, with diurnal species seeking shelter and nocturnal species emerging to hunt. The hunting ability of nocturnal reef-based predators is aided by the light of the moon. Consequently, variation in nocturnal illumination is likely to shape the timing of reproduction, larval development, and settlement for many coral reef organisms. This synthesis underscores the potential importance of trophic linkages between coral reefs and adjacent pelagic ecosystems, facilitated by the diel migrations of mesopelagic organisms and the ontogenetic migrations of coral reef larvae. Research is needed to better understand the effects of lunar cycles on life-history strategies, and the potentially disruptive effects of light pollution, turbidity, and climate-driven changes to nocturnal cloud cover. These underappreciated threats may alter patterns of nocturnal illumination that have shaped the evolutionary history of many coral reef organisms, with consequences for larval survival and population replenishment that could rival or exceed other effects arising from climate change.

Local versus site-level effects of algae on coral microbial communities.

Microbes influence ecological processes, including the dynamics and health of macro-organisms and their interactions with other species. In coral reefs, microbes mediate negative effects of algae on corals when corals are in contact with algae. However, it is unknown whether these effects extend to larger spatial scales, such as at sites with high algal densities. We investigated how local algal contact and site-level macroalgal cover influenced coral microbial communities in a field study at two islands in French Polynesia, Mo'orea and Mangareva. At 5 sites at each island, we sampled prokaryotic microbial communities (microbiomes) associated with corals, macroalgae, turf algae and water, with coral samples taken from individuals that were isolated from or in contact with turf or macroalgae. Algal contact and macroalgal cover had antagonistic effects on coral microbiome alpha and beta diversity. Additionally, coral microbiomes shifted and became more similar to macroalgal microbiomes at sites with high macroalgal cover and with algal contact, although the microbial taxa that changed varied by island. Our results indicate that coral microbiomes can be affected by algae outside of the coral's immediate vicinity, and local- and site-level effects of algae can obscure each other's effects when both scales are not considered.

Propagule redirection: habitat availability reduces colonization and increases recruitment in reef fishes.

Increased habitat availability or quality can alter production of habitat-dependent organisms in two contrasting ways: (1) by enhancing input of new colonists to the new sites (the Field-of-Dreams Hypothesis); and (2) by drawing colonists away from existing sites (the Propagule Redirection Hypothesis), and thus reducing the deleterious effects of density. We conducted a field experiment on coral reef fishes in Moorea, French Polynesia, to quantify how differing levels of habitat availability (controlling for quality) increased and/or redirected colonizing larval fish. Focal reefs without neighboring reefs received two to four times more settlers than reefs with adjacent habitat, demonstrating that increased habitat redirected larval fish. At the scale of the entire reef array, total colonization increased 1.3-fold in response to a sixfold increase in reef area (and a 2.75-fold increase in adjusted habitat availability). Thus, propagules were both increased and redirected, a result midway between the Field-of-Dreams and Propagule Redirection Hypotheses. A recruitment model using our data and field estimates of density-dependent recruitment predicts that habitat addition increases recruitment primarily by ameliorating the negative effects of competition at existing sites rather than increasing colonization at the new sites per se. Understanding long-term implications of these effects depends upon the interplay among habitat dynamics, population connectivity, colonization dynamics, and density dependence.

The vermetid gastropod Dendropoma maximum reduces coral growth and survival.

Coral reefs are one of the most diverse systems on the planet; yet, only a small fraction of coral reef species have attracted scientific study. Here, we document strong deleterious effects of an often overlooked species-the vermetid gastropod, Dendropoma maximum-on growth and survival of reef-building corals. Our surveys of vermetids on Moorea (French Polynesia) revealed a negative correlation between the density of vermetids and the per cent cover of live coral. Furthermore, the incidence of flattened coral growth forms was associated with the presence of vermetids. We transplanted and followed the fates of focal colonies of four species of corals on natural reefs where we also manipulated presence/absence of vermetids. Vermetids reduced skeletal growth of focal corals by up to 81 per cent and survival by up to 52 per cent. Susceptibility to vermetids varied among coral species, suggesting that vermetids could shift coral community composition. Our work highlights the potential importance of a poorly studied gastropod to coral dynamics.