Tim Schellekens

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.

Stage-specific predator species help each other to persist while competing for a single prey

Prey in natural communities are usually shared by many predator species. How predators coexist while competing for the same prey is one of the fundamental questions in ecology. Here, we show that competing predator species may not only coexist on a single prey but even help each other to persist if they specialize on different life history stages of the prey. By changing the prey size distribution, a predator species may in fact increase the amount of prey available for its competitor. Surprisingly, a predator may not be able to persist at all unless its competitor is also present. The competitor thus significantly increases the range of conditions for which a particular predator can persist. This “emergent facilitation” is a long-term, population-level effect that results from asymmetric increases in the rate of prey maturation and reproduction when predation relaxes competition among prey. Emergent facilitation explains observations of correlated increases of predators on small and large conspecific prey as well as concordance in their distribution patterns. Our results suggest that emergent facilitation may promote the occurrence of complex, stable, community food webs and that persistence of these communities could critically depend on diversity within predator guilds.

Coexistence of Predator and Prey in Intraguild Predation systems with Ontogenetic Niche Shifts

In basic intraguild predation (IGP) systems, predators and prey also compete for a shared resource. Theory predicts that persistence of these systems is possible when intraguild prey is superior in competition and productivity is not too high. IGP often results from ontogenetic niche shifts, in which the diet of intraguild predators changes as a result of growth in body size (life-history omnivory). As a juvenile, a life-history omnivore competes with the species that becomes its prey later in life. Competition can hence limit growth of young predators, while adult predators can suppress consumers and therewith neutralize negative effects of competition. We formulate and analyze a stage-structured model that captures both basic IGP and life-history omnivory. The model predicts increasing coexistence of predators and consumers when resource use of stage-structured predators becomes more stage specific. This coexistence depends on adult predators requiring consumer biomass for reproduction and is less likely when consumers outcompete juvenile predators, in contrast to basic IGP. Therefore, coexistence occurs when predation structures the community and competition is negligible. Consequently, equilibrium patterns over productivity resemble those of three-species food chains. Life-history omnivory thus provides a mechanism that allows intraguild predators and prey to coexist over a wide range of resource productivity.

Ontogenetic Diet Shifts Result in Niche Partitioning between Two Consumer Species Irrespective of Competitive Abilities

Tilman’s theory predicts the outcome of competition between two consumers sharing two resources on the basis of the shape of zero net‐growth isoclines (ZNGIs). In his theory, intraspecific differences in resource use are not accounted for. Here we extend this theory to include situations where organisms undergo ontogenetic diet shifts, as these characterize the life histories of many species. In a situation that without diet shifts would lead to neutral coexistence of consumer species, we investigate whether ontogenetic diet shifts lead to niche partitioning. We analyze a model describing competition for two resources between two competitors with distinctive diets over ontogeny, using copepods (showing ontogenetic diet shifts) and daphnids (not showing ontogenetic diet shifts) as appropriate representatives. We show that an ontogenetic diet shift affects the shape of the ZNGI, changing it from reflecting perfectly substitutable resources to reflecting essential resources. Furthermore, we show that resource supply determines population stage structure and stage‐dependent resource consumption in copepods and influences the competitive outcome with daphnids. In particular, we show that in itself, an ontogenetic diet shift can provide a competitive advantage if the supply of the adult resource is lower than the supply of the juvenile resource but that it always causes a disadvantage if the supply of the adult resource exceeds that of the juvenile resource.

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.