Accounting for temporal and individual variation in the estimation of Von Bertalanffy growth curves.

Growth and growth limitation are important indicators of density dependence and environmental limitation of populations. Estimating individual growth trajectories is therefore an important aspect of understanding and predicting the life history and dynamics of a population. Variation in individual growth trajectories arises due to variation in the environmental factors limiting individual growth. This environmental limitation can vary over time, between cohorts and between individuals within a cohort. For a complete and accurate understanding of individual growth in a population, it is important to include all these sources of variation. So far, statistical models only accounted for a subset of these factors or required an extensive growth history of individuals. Here, we present a novel model describing the growth curves of cohorts in a population. This model is derived from a stochastic form of the Von Bertalanffy growth equation describing individual growth. The model is specifically tailored for use on length-at-age data in which the growth trajectory of an individual is unknown and every individual is only measured once. The presented method can also be used if growth limitation differs strongly between age or length classes. We demonstrate the use of the model for length-at-age data of North Sea plaice (Pleuronectes platessa) from the last 30 years. Fitting this model to length-at-age data can provide new insights in the dynamics of the environmental factors limiting individual growth and provides a useful tool for ecological research and management.

Population-level effects of acoustic disturbance in Atlantic cod: a size-structured analysis based on energy budgets.

Anthropogenic underwater noise may negatively affect marine animals. Yet, while fishes are highly sensitive to sounds, effects of acoustic disturbances on fishes have not been extensively studied at the population level. In this study, we use a size-structured model based on energy budgets to analyse potential population-level effects of anthropogenic noise on Atlantic cod (Gadus morhua). Using the model framework, we assess the impact of four possible effect pathways of disturbance on the cod population growth rate. Through increased stress, changes in foraging and movement behaviour, and effects on the auditory system, anthropogenic noise can lead to (i) increased energy expenditure, (ii) reduced food intake, (iii) increased mortality, and (iv) reduced reproductive output. Our results show that population growth rates are particularly sensitive to changes in energy expenditure and food intake because they indirectly affect the age of maturation, survival and fecundity. Sub-lethal effects of sound exposure may thus affect populations of cod and fishes with similar life histories more than lethal effects of sound exposure. Moreover, anthropogenic noise may negatively affect populations when causing persistent increases of energy expenditure or decreases of food intake. Effects of specific acoustic pollutants on energy acquisition and expenditure should therefore be further investigated.

Estimating sensitivity of seabed habitats to disturbance by bottom trawling based on the longevity of benthic fauna.

Bottom fishing such as trawling and dredging may pose serious risks to the seabed and benthic habitats, calling for a quantitative assessment method to evaluate the impact and guide management to develop mitigation measures. We provide a method to estimate the sensitivity of benthic habitats based on the longevity composition of the invertebrate community. We hypothesize that long-lived species are more sensitive to trawling mortality due to their lower pace of life (i.e., slower growth, late maturation). We analyze data from box-core and grab samples taken from 401 stations in the English Channel and southern North Sea to estimate the habitat-specific longevity composition of the benthic invertebrate community and of specific functional groups (i.e., suspension feeders and bioturbators), and examine how bottom trawling affects the longevity biomass composition. The longevity biomass composition differed between habitats governed by differences in sediment composition (gravel and mud content) and tidal bed-shear stress. The biomass proportion of long-lived species increased with gravel content and decreased with mud content and shear stress. Bioturbators had a higher median longevity than suspension feeders. Trawling, in particular by gears that penetrate the seabed >2 cm, shifted the community toward shorter-lived species. Changes from bottom trawling were highest in habitats with many long-lived species (hence increasing with gravel content, decreasing with mud content). Benthic communities in high shear stress habitats were less affected by bottom trawling. Using these relationships, we predicted the sensitivity of the benthic community from bottom trawling impact at large spatial scale (the North Sea). We derived different benthic sensitivity metrics that provide a basis to estimate indicators of trawling impact on a continuous scale for the total community and specific functional groups. In combination with high resolution data of trawling pressure, our approach can be used to monitor and assess trawling impact and seabed status at the scale of the region or broadscale habitat and to compare the environmental impact of bottom-contacting fishing gears across fisheries.

Using marine reserves to manage impact of bottom trawl fisheries requires consideration of benthic food-web interactions.

Marine protected areas (MPAs) are widely used to protect exploited fish species as well as to conserve marine habitats and their biodiversity. They have also become a popular management tool for bottom trawl fisheries, a common fishing technique on continental shelves worldwide. The effects of bottom trawling go far beyond the impact on target species, as trawls also affect other components of the benthic ecosystem and the seabed itself. This means that for bottom trawl fisheries, MPAs can potentially be used not only to conserve target species but also to reduce impact of these side effects of the fishery. However, predicting the protective effects of MPAs is complicated because the side effects of trawling potentially alter the food-web interactions between target and non-target species. These changes in predatory and competitive interactions among fish and benthic invertebrates may have important ramifications for MPAs as tools to manage or mitigate the effects of bottom trawling. Yet, in current theory regarding the functioning of MPAs in relation to bottom trawl fisheries, such predatory and competitive interactions between species are generally not taken into account. In this study, we discuss how food-web interactions that are potentially affected by bottom trawling may alter the effectiveness of MPAs to protect (1) biodiversity and marine habitats, (2) fish populations, (3) fisheries yield, and (4) trophic structure of the community. We make the case that in order to be applicable for bottom trawl fisheries, guidelines for the implementation of MPAs must consider their potential food-web effects, at the risk of failing management.

When does fishing lead to more fish? Community consequences of bottom trawl fisheries in demersal food webs.

Bottom trawls are a globally used fishing gear that physically disturb the seabed and kill non-target organisms, including those that are food for the targeted fish species. There are indications that ensuing changes to the benthic invertebrate community may increase the availability of food and promote growth and even fisheries yield of target fish species. If and how this occurs is the subject of ongoing debate, with evidence both in favour and against. We model the effects of trawling on a simple ecosystem of benthivorous fish and two food populations (benthos), susceptible and resistant to trawling. We show that the ecosystem response to trawling depends on whether the abundance of benthos is top-down or bottom-up controlled. Fishing may result in higher fish abundance, higher (maximum sustainable) yield and increased persistence of fish when the benthos which is the best-quality fish food is also more resistant to trawling. These positive effects occur in bottom-up controlled systems and systems with limited impact of fish feeding on benthos, resembling bottom-up control. Fishing leads to lower yields and fish persistence in all configurations where susceptible benthos are more profitable prey. Our results highlight the importance of mechanistic ecosystem knowledge as a requirement for successful management.

Coexistence of two stage-structured intraguild predators.

An organism can be defined as omnivorous if it feeds on more than one trophic level. Omnivory is present in many ecosystems and multiple omnivorous species can coexist in the same ecosystem. How coexisting omnivores are able to avoid competitive exclusion is very much an open question. In this paper we analyze a model of a community consisting of two omnivorous predators and a basal resource. The population of both predators is explicitly structured into juveniles and adults, of which juveniles only feed on basal resource and adults feed on a varied proportion of basal resource and juveniles of the other population. We thereby separate the omnivorous roles (competitor for basal resource and predator of competitors) over life history. We show in this study that persistence of multiple omnivorous predators is possible when predators differ in adult diets. In this case, coexistence occurs because community dynamics force one of the model species to act as a predator and the other to act as a consumer. We conclude that separation of omnivorous roles over life history not only offers an explanation on why systems with omnivory can persist, but also how multiple omnivores can coexist at the same trophic levels of those systems.

Interspecific resource competition effects on fisheries revenue.

In many fisheries multiple species are simultaneously caught while stock assessments and fishing quota are defined at species level. Yet species caught together often share habitat and resources, resulting in interspecific resource competition. The consequences of resource competition on population dynamics and revenue of simultaneously harvested species has received little attention due to the historical single stock approach in fisheries management. Here we present the results of a modelling study on the interaction between resource competition of sole (Solea solea) and slaice (Pleuronectus platessa) and simultaneous harvesting of these species, using a stage-structured population model. Three resources were included of which one is shared with a varied competition intensity. We find that plaice is the better competitor of the two species and adult plaice are more abundant than adult sole. When competition is high sole population biomass increases with increasing fishing effort prior to plaice extinction. As a result of this increase in the sole population, the revenue of the stocks combined as function of effort becomes bimodal with increasing resource competition. When considering a single stock quota for sole, its recovery with increasing effort may result in even more fishing effort that would drive the plaice population to extinction. When sole and plaice compete for resources the highest revenue is obtained at effort levels at which plaice is extinct. Ignoring resource competition promotes overfishing due to increasing stock of one species prior to extinction of the other species. Consequently, efforts to mitigate the decline in one species will not be effective if increased stock in the other species leads to increased quota. If a species is to be protected against extinction, management should not only be directed at this one species, but all species that compete with it for resource as well.

Complete compensation in Daphnia fecundity and stage-specific biomass in response to size-independent mortality.

1. Recent theory suggests that compensation or even overcompensation in stage-specific biomass can arise in response to increased mortality. Which stage that will show compensation depends on whether maturation or reproduction is the more limiting process in the population. Size-structured theory also provides a strong link between the type of regulation and the expected population dynamics as both depend on size/stage-specific competitive ability. 2. We imposed a size-independent mortality on a consumer-resource system with Daphnia pulex feeding on Scenedesmus obtusiusculus to asses the compensatory responses in Daphnia populations. We also extended an existing stage-structured biomass model by including several juvenile stages to test whether this extension affected the qualitative results of the existing model. 3. We found complete compensation in juvenile biomass and total population fecundity in response to harvesting. The compensation in fecundity was caused by both a higher proportion of fecund females and a larger clutch size under increased mortality. We did not detect any difference in resource levels between treatments. 4. The model results showed that both stages of juveniles have to be superior to adults in terms of resource competition for the compensatory response to take place in juvenile biomass. 5. The results are all in correspondence with that the regulating process within the population was reproduction. From this, we also conclude that juveniles were superior competitors to adults, which has implications for population dynamics and the kind of cohort cycles seen in Daphnia populations. 6. The compensatory responses demonstrated in this experiment have major implications for community dynamics and are potentially present in any organisms with food-dependent growth or development.

Invasion success depends on invader body size in a size-structured mixed predation-competition community.

1. The size of an individual is an important determinant of its trophic position and the type of interactions it engages in with other heterospecific and conspecific individuals. Consequently an individual's ecological role in a community changes with its body size over ontogeny, leading to that trophic interactions between individuals are a size-dependent and ontogenetically variable mixture of competition and predation. 2. Because differently sized individuals thus experience different biotic environments, invasion success may be determined by the body size of the invaders. Invasion outcome may also depend on the productivity of the system as productivity influences the biotic environment. 3. In a laboratory experiment with two poeciliid fishes the body size of the invading individuals and the daily amount of food supplied were manipulated. 4. Large invaders established persistent populations and drove the resident population to extinction in 10 out of 12 cases, while small invaders failed in 10 out of 12 trials. Stable coexistence was virtually absent. Invasion outcome was independent of productivity. 5. Further analyses suggest that small invaders experienced a competitive recruitment bottleneck imposed on them by the resident population. In contrast, large invaders preyed on the juveniles of the resident population. This predation allowed the large invaders to establish successfully by decreasing the resident population densities and thus breaking the bottleneck. 6. The results strongly suggest that the size distribution of invaders affects their ability to invade, an implication so far neglected in life-history omnivory systems. The findings are further in agreement with predictions of life-history omnivory theory, that size-structured interactions demote coexistence along a productivity gradient.

Simplifying a physiologically structured population model to a stage-structured biomass model.

We formulate and analyze an archetypal consumer-resource model in terms of ordinary differential equations that consistently translates individual life history processes, in particular food-dependent growth in body size and stage-specific differences between juveniles and adults in resource use and mortality, to the population level. This stage-structured model is derived as an approximation to a physiologically structured population model, which accounts for a complete size-distribution of the consumer population and which is based on assumptions about the energy budget and size-dependent life history of individual consumers. The approximation ensures that under equilibrium conditions predictions of both models are completely identical. In addition we find that under non-equilibrium conditions the stage-structured model gives rise to dynamics that closely approximate the dynamics exhibited by the size-structured model, as long as adult consumers are superior foragers than juveniles with a higher mass-specific ingestion rate. When the mass-specific intake rate of juvenile consumers is higher, the size-structured model exhibits single-generation cycles, in which a single cohort of consumers dominates population dynamics throughout its life time and the population composition varies over time between a dominance by juveniles and adults, respectively. The stage-structured model does not capture these dynamics because it incorporates a distributed time delay between the birth and maturation of an individual organism in contrast to the size-structured model, in which maturation is a discrete event in individual life history. We investigate model dynamics with both semi-chemostat and logistic resource growth.

Size-dependent mortality induces life-history changes mediated through population dynamical feedbacks.

The majority of taxa grow significantly during life history, which often leads to individuals of the same species having different ecological roles, depending on their size or life stage. One aspect of life history that changes during ontogeny is mortality. When individual growth and development are resource dependent, changes in mortality can affect the outcome of size-dependent intraspecific resource competition, in turn affecting both life history and population dynamics. We study the outcome of varying size-dependent mortality on two life-history types, one that feeds on the same resource throughout life history and another that can alternatively cannibalize smaller conspecifics. Compensatory responses in the life history dampen the effect of certain types of size-dependent mortality, while other types of mortality lead to dramatic changes in life history and population dynamics, including population (de-)stabilization, and the growth of cannibalistic giants. These responses differ strongly among the two life-history types. Our analysis provides a mechanistic understanding of the population-level effects that come about through the interaction between individual growth and size-dependent mortality, mediated by resource dependence in individual vital rates.

Food-dependent growth leads to overcompensation in stage-specific biomass when mortality increases: the influence of maturation versus reproduction regulation.

We analyze a stage-structured biomass model for size-structured consumer-resource interactions. Maturation of juvenile consumers is modeled with a food-dependent function that consistently translates individual-level assumptions about growth in body size to the population level. Furthermore, the model accounts for stage-specific differences in resource use and mortality between juvenile and adult consumers. Without such differences, the model reduces to the Yodzis and Innes (1992) bioenergetics model, for which we show that model equilibria are characterized by a symmetry property that reproduction and maturation are equally limited by food density. As a consequence, biomass production rate exactly equals loss rate through maintenance and mortality in each consumer stage. Stage-specific differences break up this symmetry and turn specific stages into net producers and others into net losers of biomass. As a consequence, the population in equilibrium can be regulated in two distinct ways: either through total population reproduction or through total population maturation as limiting process. In the case of reproduction regulation, increases in mortality may lead to an increase of juvenile biomass. In the case of maturation regulation, increases in mortality may increase adult biomass. This overcompensation in biomass occurs with increases in both stage-independent and stage-specific mortality, even when the latter targets the stage exhibiting overcompensation.

Population dynamical consequences of gregariousness in a size-structured consumer-resource interaction.

Many animal species live in groups. Group living may increase exploitation competition within the group, and variation among groups in intra-group competition intensity could induce life-history variability among groups. Models of physiologically structured populations generally predict single generation cycles, driven by exploitation competition within and between generations. We expect that life-history variability and habitat heterogeneity induced by group living may affect such competition-driven population dynamics. In this study, we vary the gregariousness (the tendency to aggregate in groups) of a size-structured consumer population in a spatially explicit environment. The consumer has limited mobility, and moves according to a probabilistic movement process. We study the effects on the population dynamics, as mediated through the resource and the life-history of the consumer. We find that high gregariousness leads to large spatial resource variation, and highly variable individual life-history, resulting in highly stochastic population dynamics. At reduced gregariousness, life-history of consumers synchronizes, habitat heterogeneity is reduced, and single generation cycles appear. We expect this pattern to occur for any group living organism with limited mobility. Our results indicate that constraints set by population dynamical feedback may be an important aspect in understanding group living in nature.

Bistability and an Allee effect as emergent consequences of stage-specific predation.

The Allee effect, a reduction of individual fitness at low population density that can lead to sudden and unannounced extinctions, has been shown to come about through a number of mechanisms, usually associated with group behavior or mate search. Recent papers show that it may arise through size-selective predation, without explicit assumptions relating individual fitness to population density. It arises from the shift that a predator induces in the population stage distribution of its prey. We study the parameter conditions that lead to such an emergent Allee effect. The emergent Allee effect occurs under fairly broad conditions. We show that stage-specific predation can also induce bistability between alternative states where both prey and predator are present. A perturbation analysis on the equilibria shows that all equilibria are highly robust to changes in predator density. Our work shows that when size-specific interactions are taken into account, bistabilities and catastrophic collapses are possible even in purely exploitative food webs, which has substantial implications for questions related to food web theory and conservation issues.