The relative importance of metabolic rate and body size to space use behavior in aquatic invertebrates.

Elucidating the underlying mechanisms behind variations of animal space and resource use is crucial to pinpoint relevant ecological phenomena. Organism's traits related to its energy requirements might be central in explaining behavioral variation, as the ultimate goal of a forager is to fulfill its energy requirements. However, it has remained poorly understood how energy requirements and behavioral patterns are functionally connected. Here we aimed to assess how body mass and standard metabolic rate (SMR) influence behavioral patterns in terms of cumulative space use and time spent in an experimental patchy environment, both within species and among individuals irrespective of species identity. We measured the behavioral patterns and SMR of two invertebrate species, that is, amphipod Gammarus insensibilis, and isopod Lekanesphaera monodi, individually across a range of body masses. We found that species of G. insensibilis have higher SMR level, in addition to cumulatively exploring a larger space than L. monodi. Cumulative space use scaled allometrically with body mass, and it scaled isometrically with SMR in both species. While time spent similarly in both species was characterized by negative body mass and SMR dependence, it was observed that L. monodi individuals tended to stay longer in resource patches compared to G. insensibilis individuals. Our results further showed that within species, body mass and metabolic rate explained a similar amount of variation in behavior modes. However, among individuals, regardless of species identity, SMR had stronger predictive power for behavioral modes compared to body mass. This suggests that SMR might offer a more generalized and holistic description of behavioral patterns that extend beyond species identity. Our study on the metabolic and body mass scaling of space and resource use behavior sheds light on higher-order ecological processes such as species' competitive coexistence along the spatial and trophic dimensions.

Metabolic rate and climate change across latitudes: evidence of mass-dependent responses in aquatic amphipods.

Predictions of individual responses to climate change are often based on the assumption that temperature affects the metabolism of individuals independently of their body mass. However, empirical evidence indicates that interactive effects exist. Here, we investigated the response of individual standard metabolic rate (SMR) to annual temperature range and forecasted temperature rises of 0.6-1.2°C above the current maxima, under the conservative climate change scenario IPCC RCP2.6. As a model organism, we used the amphipod Gammarus insensibilis, collected across latitudes along the western coast of the Adriatic Sea down to the southernmost limit of the species' distributional range, with individuals varying in body mass (0.4-13.57 mg). Overall, we found that the effect of temperature on SMR is mass dependent. Within the annual temperature range, the mass-specific SMR of small/young individuals increased with temperature at a greater rate (activation energy: E=0.48 eV) than large/old individuals (E=0.29 eV), with a higher metabolic level for high-latitude than low-latitude populations. However, under the forecasted climate conditions, the mass-specific SMR of large individuals responded differently across latitudes. Unlike the higher-latitude population, whose mass-specific SMR increased in response to the forecasted climate change across all size classes, in the lower-latitude populations, this increase was not seen in large individuals. The larger/older conspecifics at lower latitudes could therefore be the first to experience the negative impacts of warming on metabolism-related processes. Although the ecological collapse of such a basic trophic level (aquatic amphipods) owing to climate change would have profound consequences for population ecology, the risk is significantly mitigated by phenotypic and genotypic adaptation.

The size dependency of foraging behaviour: an empirical test performed on aquatic amphipods.

The behavioural choices made by foragers regarding the use of resource patches have a direct influence on the energy balance of the individual. Given that several individual traits linked to the acquisition of spatially distributed resources increase with body size (e.g., energy requirements, resource ingestion rates, and movement capacity), it is reasonable to expect size dependencies in overall foraging behaviour. In this study, we tested how body size influences the number, duration, and frequency of foraging episodes in heterogeneous resource patches. To this end, we performed microcosm experiments using the aquatic amphipod Gammarus insensibilis as a model organism. An experimental maze was used to simulate a habitat characterised by resource-rich, resource-poor, and empty patches under controlled conditions. The patch use behaviour of 40 differently sized specimens foraging alone in the experimental maze was monitored via an advanced camera setup. Overall, we observed that individual body size exerted a major influence on the use of resource patches over time. Larger individuals had stronger preference for the resource-rich patches initially and visited them more frequently than smaller individuals, but for shorter periods of time. However, larger individuals subsequently decreased their use of resource-rich patches in favour of resource-poor patches, while smaller individuals continued to prefer resource-rich patches for the whole experimental time. With body size being a key organismal trait, our observations support the general understanding of foraging behaviours related to preference, patch use, and abandonment.

Modelling spatial and temporal patterns in bioturbator effects on sediment resuspension: A biophysical metabolic approach.

Tidal flats are biogeomorphic landscapes, shaped by physical forces and interaction with benthic biota. We used a metabolic approach to assess the overarching effect of bioturbators on tidal landscapes. The benthic bivalve common cockle (Cerastoderma edule) was used as model organism. The effect of C. edule on sediment resuspension was approximated as a function of the overall population metabolic rate per unit of area. We combined i) laboratory observations on how C. edule affect sediment resuspension along gradients of bioturbation activity, sediment cohesiveness and hydrodynamic force with ii) spatial data on the natural distribution of intertidal C. edule populations. This allowed us to build an integrated model of the C. edule effect on sediment resuspension along the tidal gradient. Owing to the temperature dependence of metabolic rate, the model also accounted for seasonal variation in bioturbators activity. Laboratory experiments indicated that sediment resuspension is positively related to the metabolic rate of the C. edule population especially in cohesive sediments. Based on this observation, we predicted a clear spatial and seasonal pattern in the relative importance of C. edule contribution to sediment resuspension along a tidal transect. At lower elevations, our model indicates that hydrodynamics overrules biotic effects; at higher elevations, inter-tidal hydrodynamics should be too low to suspend bioturbated sediments. The influence of C. edule on sediment resuspension is expected to be maximal at the intermediate elevation of a mudflat, owing to the combination of moderate hydrodynamic stress and high bioturbator activity. Also, bio-mediated sediment resuspension is predicted to be particularly high in the warm season. Research into metabolic dependency of bio-mediated sediment resuspension may help to place phenomenological observations in the broader framework of metabolic theories in ecology and to formulate general expectations on the coastal ecosystem functioning.

A new approach to assessing the space use behavior of macroinvertebrates by automated video tracking.

Individual space and resource use are central issues in ecology and conservation. Recent technological advances such as automated tracking techniques are boosting ecological research in this field. However, the development of a robust method to track space and resource use is still challenging for at least one important ecosystem component: motile aquatic macroinvertebrates. The challenges are mostly related to the small body size and rapid movement of many macroinvertebrate species and to light scattering and wave signal interference in aquatic habitats.We developed a video tracking method designed to reliably assess space use behavior among individual aquatic macroinvertebrates under laboratory (microcosm) conditions. The approach involves the use of experimental apparatus integrating a near infrared backlight source, a Plexiglas multi-patch maze, multiple infrared cameras, and automated video analysis. It allows detection of the position of fast-moving (~ 3 cm/s) and translucent individuals of small size (~ 5 mm in length, ~1 mg in dry weight) on simulated resource patches distributed over an experimental microcosm (0.08 m2).To illustrate the adequacy of the proposed method, we present a case study regarding the size dependency of space use behavior in the model organism Gammarus insensibilis, focusing on individual patch selection, giving-up times, and cumulative space used.In the case study, primary data were collected on individual body size and individual locomotory behavior, for example, mean speed, acceleration, and step length. Individual entrance and departure times were recorded for each simulated resource patch in the experimental maze. Individual giving-up times were found to be characterized by negative size dependency, with patch departure occurring sooner in larger individuals than smaller ones, and individual cumulative space used (treated as the overall surface area of resource patches that individuals visited) was found to scale positively with body size.This approach to studying space use behavior can deepen our understanding of species coexistence, yielding insights into mechanistic models on larger spatial scales, for example, home range, with implications for ecological and evolutionary processes, as well as for the management and conservation of populations and ecosystems. Despite being specifically developed for aquatic macroinvertebrates, this method can also be applied to other small aquatic organisms such as juvenile fish and amphibians.

Size dependency of patch departure behavior: evidence from granivorous rodents.

Individual size is a major determinant of mobile organisms' ecology and behavior. This study aims to explore whether allometric scaling principles can provide an underlying framework for general patterns of resource patch use. To this end, we used giving-up densities (GUDs), that is, the amount of resources remaining in a patch after a forager has quit feeding, as a comparative measure of the amount of resources exploited by a forager of any given size. We specifically tested the hypothesis that size-dependent responses to both internal (energy requirement) and external (risk management) forces may have an effect on GUDs. We addressed this topic by conducting an extensive meta-analysis of published data on granivorous rodents, including 292 GUD measurements reported in 25 papers. The data set includes data on 22 granivorous rodent species belonging to three taxonomic suborders (Castorimorpha, Myomorpha, and Sciuromorpha) and spans three habitat types (desert, grassland, and forest). The observations refer to both patches subject to predation risk and safe patches. Pooling all data, we observed positive allometric scaling of GUDs with average forager size (scaling exponent = 0.45), which explained 15% of overall variance in individual GUDs. Perceived predation risk during foraging led to an increase in GUDs independently of forager size and taxonomy and of habitat type, which explained an additional 12% of overall GUD variance. The size scaling exponent of GUDs is positive across habitat types and taxonomic suborders of rodents. Some variation was observed, however. The scaling coefficients in grassland and forest habitat types were significantly higher than in the desert habitat type. In addition, Sciuromorpha and Myomorpha exhibited a more pronounced size scaling of GUDs than Castorimorpha. This suggests that different adaptive behaviors may be used in different contexts and/or from different foragers. With body size being a fundamental ecological descriptor, research into size scaling of GUDs may help to place patch-use observations in a broader allometric framework.

A process based model of cohesive sediment resuspension under bioturbators' influence.

Macrozoobenthos may affect sediment stability and erodibility via their bioturbating activities, thereby impacting both the short- and long-term development of coastal morphology. Process-based models accounting for the effect of bioturbation are needed for the modelling of erosion dynamics. With this work, we explore whether the fundamental allometric principles of metabolic activity scaling with individual and population size may provide a framework to derive general patterns of bioturbation effect on cohesive sediment resuspension. Experimental flumes were used to test this scaling approach across different species of marine, soft-sediment bioturbators. The collected dataset encompasses a range of bioturbator functional diversity, individual densities, body sizes and overall population metabolic rates. Measurements were collected across a range of hydrodynamic stress from 0.02 to 0.25 Pa. Overall, we observed that bioturbators are able to slightly reduce the sediment resuspension at low hydrodynamic stress, whereas they noticeably enhance it at higher levels of stress. Along the whole hydrodynamic stress gradient, the quantitative effect of bioturbators on sediment resuspension can be efficiently described by the overall metabolic rate of the bioturbating benthic communities, with significant variations across the bioturbators' taxonomic and functional diversity. One of the tested species (the gallery-builder Polychaeta Hediste diversicolor) had an effect that was partially deviating from the general trend, being able to markedly reduce sediment resuspension at low hydrodynamic stress compared to other species. By combining bioturbators' influence with hydrodynamic force, we were able to produce a process-based model of biota-mediated sediment resuspension.

Revisiting GUD: An empirical test of the size-dependency of patch departure behaviour.

Behaviour related to patch resource exploitation is a major determinant of individual fitness. Assuming the size-dependency of patch departure behaviour, model-based approaches have shown size-mediated coexistence in systems of competing species. However, experimental evidence for the influence of body size on patch use behaviour is scarce. In this study, we explore whether allometric principles provide an underlying framework for interspecific patterns of resource use. To this end, we propose a meso-cosm approach using three species of gastropods differing in size as a model system and 32P radio-isotopic techniques as a measure of resource use. Foragers of different size were placed in an artificial patch, provided with a limited amount of labelled resource and let them free to move as resources decrease and scarcity is sensed. We investigated the extent to which individual body size affects the exploitation of resources by examining Giving Up Density (GUD), Giving Up Time (GUT), resource absorption rate and exploitation efficiency as components of individual exploitation behaviour. To compare positive, constant and negative individual size scaling of population energy requirements, experimental trials with an equal numbers and equal biomass of differently sized foragers were carried out, and an experimental trial with equal metabolic requirements was simulated. We observed clear size dependency in the patch departure behaviour of the experimental organisms. Even under conditions of equivalent overall population energy requirements, larger foragers decided to leave the resource patch earlier and at a higher density of resources than smaller ones. Smaller foragers were able to prolong their presence and make more use of the resources, resulting in an inverse body-size scaling of resource exploitation efficiency.

The combined influence of body size and density on cohesive sediment resuspension by bioturbators.

We propose an empirical framework to scale the effects of bioturbation on sediment resuspension to population bioturbation activity, approximated as population metabolic rate. Individual metabolic rates have been estimated as functions of body size and extrapolated to population level. We used experimental flumes to test this approach across different types of marine, soft-sediment bioturbators. We observed that a large part of the variance in biota-mediated sediment resuspension can be explained by a positive relationship with population metabolic rate. Other mechanisms can strongly influence the outcome, such as bioturbation of deep sediment strata, biotic interactions with hydrodynamic stress and overlapping areas of influence must be further investigated. By relating the biota-mediated changes in resuspended sediment to metabolism, we can place our observations within the broader context of the metabolic theory of ecology and to formulate general expectations about changes in biota-mediated sediment resuspension in response to changes in population structure and climate change.

Analysis of macrobenthic assemblages and ecological health of Yellow River Delta, China, using AMBI & M-AMBI assessment method.

Yellow River delta (YRD) is a typical example of a valuable coastal ecosystem that is under increasing anthropogenic threat in China. To understand the current health status of this region, three surveys in 2011 for the abiotic conditions and macrobenthic assemblages were performed. The concentration of trace metals were relatively low in the sediment at all sampling stations representing a good sediment quality. A total of 159 macrobenthic species were identified during the three surveys. ABC curves showed that the macrobenthic fauna at 8 sampling stations suffered disturbances from human activities. M-AMBI index indicates that the benthic ecological quality of YRD is currently still not in a good condition. Five trace metals, water temperature and depth were the main environmental variables affecting the distribution pattern of macrobenthic assemblages. Community succession has occurred over the past 60years, as evidenced by changes of species composition, key species, distribution pattern and range.

A mixed modeling approach to predict the effect of environmental modification on species distributions.

Human infrastructures can modify ecosystems, thereby affecting the occurrence and spatial distribution of organisms, as well as ecosystem functionality. Sustainable development requires the ability to predict responses of species to anthropogenic pressures. We investigated the large scale, long term effect of important human alterations of benthic habitats with an integrated approach combining engineering and ecological modelling. We focused our analysis on the Oosterschelde basin (The Netherlands), which was partially embanked by a storm surge barrier (Oosterscheldekering, 1986). We made use of 1) a prognostic (numerical) environmental (hydrodynamic) model and 2) a novel application of quantile regression to Species Distribution Modeling (SDM) to simulate both the realized and potential (habitat suitability) abundance of four macrozoobenthic species: Scoloplos armiger, Peringia ulvae, Cerastoderma edule and Lanice conchilega. The analysis shows that part of the fluctuations in macrozoobenthic biomass stocks during the last decades is related to the effect of the coastal defense infrastructures on the basin morphology and hydrodynamics. The methodological framework we propose is particularly suitable for the analysis of large abundance datasets combined with high-resolution environmental data. Our analysis provides useful information on future changes in ecosystem functionality induced by human activities.