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Terrestrial runoff into aquatic ecosystems may have both stimulatory and inhibitory effects, due to nutrient subsidies and increased light attenuation. To disentangle the effects of runoff on microbenthos, we added soil to coastal mesocosms and manipulated substrate depth. To test if fish interacted with runoff effects, we manipulated fish presence. Soil decreased microphytobenthic chlorophyll-a per area and per carbon (C) unit, increased microbenthic phosphorous (P), and reduced microbenthic nitrogen (N) content. Depth had a strong effect on the microbenthos, with shallow substrates exhibiting greater microbenthic net ecosystem production, gross primary production, and community respiration than deep substrates. Over time, micobenthic algae compensated for deeper substrate depth through increased chlorophyll-a synthesis, but despite algal shade compensation, the soil treatment still appeared to reduce the depth where microbenthos switched from net autotrophy to net heterotrophy. Fish interacted with soil in affecting microbenthic nutrient composition. Fish presence reduced microbenthic C/P ratios only in the no soil treatment, probably since soil nutrients masked the positive effects of fish excreta on microbenthos. Soil reduced microbenthic N/P ratios only in the absence of fish. Our study demonstrates the importance of light for the composition and productivity of microbenthos but finds little evidence for positive runoff subsidy effects.

Heavy rainfall events causing significant terrestrial runoff into coastal marine ecosystems are predicted to become more frequent with climate change in the Mediterranean. To simulate the effects of soil runoff on the pelagic food web of an oligotrophic Mediterranean coastal lagoon, we crossed soil extract addition (increasing nutrient availability and turbidity) and fish presence in a full factorial design to coastal mesocosms containing a natural pelagic community. Soil extract addition increased both bacteria and phytoplankton biomass. Diatoms however profited most from soil extract addition, especially in the absence of fish. In contrast zooplankton and fish did not profit from soil extract addition. Furthermore, our data indicate that nutrients (instead of light or carbon) limited basal production. Presumed changes in carbon availability are relatively unimportant to primary and secondary production in strongly nutrient limited systems like the Thau Lagoon. We conclude that in shallow Mediterranean coastal ecosystems, heavy rainfall events causing soil runoff will (1) increase the relative abundance of phytoplankton in relation to bacteria and zooplankton, especially in the absence of fish (2) not lead to higher biomass of zooplankton and fish, possibly due to the brevity of the phytoplankton bloom and the slow biomass response of higher trophic levels.

Size-dependent differences between individuals in size-structured organisms have fundamental effect on population and community dynamics. Intraguild predation (IGP) is one specifically interesting constellation that often arises when two size-structured populations interact. Ontogenetic bottlenecks that determine population size-structure are affected by both population intrinsic as well as population extrinsic factors, and are therefore context-dependent. Surprisingly, size-structured IGP systems have mainly been investigated theoretically and especially long-term empirical studies are widely lacking. In this thesis I investigate empirically how habitat complexity, interaction strength, and stage-specific resource availabilities affect population processes and their effects on the dynamics of a size-structured IGP system. I conducted multi-generation experiments in a size-structured IGP system, with the Least Killifish (Heterandria formosa) as IG prey and the Common Guppy (Poecilia reticulata) as IG predator. With no alternative resource next to the shared resource, IG predator and IG prey could not coexist. Weak interactions only increased IG prey and IG predator persistence times and observed exclusion patterns depended on habitat complexity. An alternative resource for either the juvenile IG predator or the juvenile IG prey on the other hand promoted coexistence. However, this coexistence was context-dependent. Ontogenetic bottlenecks played a central role in the dynamics of the size-structured IGP system in general. In the final study I show that an ontogenetic bottleneck can, through changes in stage-specific resource availabilities, be affected in a way that leads to increased trophic transfer efficiency with potential effects on higher trophic levels.

Overall, the results emphasize importance of the broader context in which size-structured communities are embedded. Especially, when managing natural communities it is important to account for the combined effects of size-structure, stage-specific resource availabilities, and habitat structure. Specifically, when managing species that connect habitats or ecosystems all life-stages’ environmental conditions must be consider in order to ensure strong predictive power of tools used for ecosystem management planning.

Patterns in biomass production are determined by resource input (productivity) and trophic transfer efficiency. At fixed resource input, variation in consumer biomass production has been related to food quality, metabolic type and diversity among species. In contrast, intraspecific variation in individual body size because of ontogenetic development, which characterizes the overwhelming majority of taxa, has been largely neglected. Here we show experimentally in a long-term multigenerational study that reallocating constant resource input in a two-stage consumer system from an equal resource delivery to juveniles and adults to an adult-biased resource delivery is sufficient to cause more than a doubling of total consumer biomass. We discuss how such changes in consumer stage-specific resource allocation affect the likelihood for alternative stable states in harvested populations as a consequence of stage-specific overcompensation in consumer biomass and thereby the risk of catastrophic collapses in exploited populations.

In life history omnivore (IGP) systems coexistence between omnivore and consumer at high productivity has only been demonstrated when the omnivore undergoes a complete ontogenetic niche shift at or before maturity from feeding on the shared resource to feeding on the consumer. Here we investigate the effects of an exclusive resource for juvenile consumers on coexistence between omnivore and consumer. We demonstrate that an alternative resource for juvenile consumers allows for coexistence between omnivore and consumer species even when the adult omnivore feeds on the shared resource to a substantial extent. Coexistence is promoted by a strong niche separation in the consumer and when the productivity of the shared resource is high relative to the juvenile consumer exclusive resource. At high shared resource productivity coexistence is promoted by either a low or a high niche separation in the omnivore. In general our results suggest that for coexistence to occur at high productivities a strong life-history separation in resource use is necessary in either the consumer or the omnivore. Strong life-history separation in the omnivore results in predation driven coexistence, while strong life-history separation in the consumer results in competitive coexistence. 

Coexistence in size-structured intraguild predation systems is limited, and is predicted to occur when the IG predator exhibits a complete ontogenetic niche shift, has a similar asymptotic size as the IG prey, or has access to an exclusive resource. Here we experimentally test the effect of a juvenile IG predator exclusive resource by mimicking an ontogenetic habitat shift in the IG predator that switched either to a more or a less productive habitat at maturity. We show that coexistence depends on relative habitat productivities when refuges are present and cannibalism in the IG predator is low, but does not so when refuges are absent and cannibalism in the IG predator is substantial. Overall compared to previous experimental studies without an ontogenetic habitat shift, we how that an exclusive resource for juvenile IG predators promotes coexistence with context-dependent differences in the manifestation of this effect. 

Complex habitats and thereby weaker predator-prey interactions have been suggested to promote coexistence between predator and prey in intraguild predation (IGP) systems. For a size-structured IGP system spatial refuges have been shown to weaken interactions but not to promote coexistence. Spatial refuges however also affect the spatial distribution of small and large individuals. Here we report the results of a multi-generation laboratory experiment where we manipulated interaction strength by using the same IG predator (Common guppy, Poecilia reticulata) but a population with a different life-history evolution and lower predation voracity. Resident IG prey (Least Killifish, Heterandria formosa) were invaded by large or small IG predators, invasion success was recorded. Compared to the invasion by more voracious IG predator individuals, weaker predation per se (no refuges) did not affect invasion success but did increase IG prey and IG predator persistence times. Compared to the invasion by more voracious IG predator individuals in the presence of refuges, weaker predation per se (no refuges) resulted in similar persistence times but different invasion success.  We conclude that the effect on community dynamics depends on the context in which weak interactions are realized. Both spatial refuges and life-history differences affected predation strength and competitive relationships quantitatively but only when spatial refuges were present was this quantitative change coupled to qualitative changes in species interactions. Though under stable environmental conditions in our experiment coexistence did not occur we argue that in temporarily or spatially variable systems weak interactions have the potential to promote coexistence by prolonging IG predator and IG prey persistence times.

Size-dependent interactions and habitat complexity have been identified as important factors affecting the persistence of intraguild predation (IGP) systems. Habitat complexity has been suggested to promote intraguild (IG) prey and intraguild predator coexistence through weakening trophic interactions particularly the predation link. Here, we experimentally investigate the effects of habitat complexity on coexistence and invasion success of differently sized IG-predators in a size-structured IGP system consisting of the IG-predator Poecilia reticulata and a resident Heterandria formosa IG-prey population. The experiments included medium-long and long-term invasion experiments, predator-prey experiments and competition experiments to elucidate the mechanisms underlying the effect of prey refuges. Habitat complexity did not promote the coexistence of IG-predator and IG-prey, although the predation link was substantially weakened. However, the presence of habitat structure affected the invasion success of large IG-predators negatively and the invasion success of small IG-predators positively. The effect of refuges on size-dependent invasion success could be related to a major decrease in the IG-predator's capture rate and a shift in the size distribution of IG-predator juveniles. In summary, habitat complexity had two main effects: (i) the predation link was diminished, resulting in a more competition driven system and (ii) the overall competitive abilities of the two species were equalized, but coexistence was not promoted. Our results suggest that in a size-structured IGP system, individual level mechanisms may gain in importance over species level mechanisms in the presence of habitat complexity.

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.