Eva Wahlström

Size-specific interactions in lake systems : Predator gape limitation and prey growth rate and mortality

To study the effects of different predators on the behavior and dynamics of their prey, we compared the performance of perch (Perca fluviatilis) and the abundance of their prey resource in four lakes with two different types of piscivory. In two of the lakes, perch were the only fish species present and piscivory was restricted to cannibalism. In the other two lakes, perch co-occurred with piscivorous pike (Esox lucius). Pike grow to larger sizes and can capture larger prey than piscivorous perch. Therefore, perch reach a size refuge earlier in the lakes without pike. Perch in the lakes lacking pike used both the epilimnion and hypolimnion habitats of the lakes, and a high proportion of the perch occurred below the thermocline. In contrast, perch in the lakes with pike were mainly captured along the littoral zone and were never captured below the thermocline. This difference in habitat use was reflected in different diets of perch in the different lakes. Perch in the lakes with pike were also less abundant than those in the lakes lacking pike and the growth rates of individual perch and the densities of predator-sensitive prey (pelagic and littoral) were higher, indicating that the presence of pike resulted in decreased intraspecific competition in perch. Evidence for a behavioral response to predation risk was found in 1+ (1 yr old) perch, which was suggested to be due to predation risk from both larger perch and pike. It is hypothesized that the use of the cold-water hypolimnion habitat by perch in the lakes lacking pike was a response of perch to reduce metabolic demands under conditions of severe resource limitation. Differences in gape limitation between predators species are suggested to have major effects on size-dependent mortalities and growth rates in prey and thereby on overall community dynamics. Evidence for this latter effect was observed in differences in zoo- plankton size structure between the lakes with and without pike.

CANNIBALISM AND COMPETITION IN EURASIAN PERCH: POPULATION DYNAMICS OF AN ONTOGENETIC OMNIVORE

In many cannibalistic populations, cannibals share resources with their victims, leading to a size-dependent mixture of cannibalistic and competitive interactions. We analyze the impacts of such interactions on the population dynamics of Eurasian perch (Percafluviatilis) by considering effects of intercohort competition, habitat heterogeneity, habitat selection, and energy gain made by cannibals. Over a six-year period, we measured mortality and recruitment patterns, individual growth, body condition, resource levels, diets, and habitat use as functions of density for an allopatric perch population in a low-productivity lake. During the course of the study, two major die-offs took place, selectively affecting larger cannibalistic individuals, followed by several years of successful recruitment of young fish. Habitat use of perch ?2 yr old was density dependent, and these fish used only the inshore region at low densities. The appearance of young fish followed the die-offs of cannibalistic perch and their subsequent absence from the offshore area, both of which decreased cannibalism on pelagic recruits. Whereas die-offs of larger perch could not be related to competition with young-of-the-year (YOY) perch, evidence for a competitive impact of YOY perch on Age-I perch was present due to substantial food overlap. A strong depression in pelagic zooplankton was observed during summer in years with strong re- cruitment, which resulted in reduced consumption of zooplankton, slow growth, and reduced body condition in Age-1 perch and suggested high mortality of Age-1 perch in autumn. Age-I perch did not appear to profit substantially from cannibalism on YOY perch because of the short time period that they could efficiently prey on YOY perch. The few larger perch that survived the die-offs gained substantial energy from cannibalism in years with strong recruitments, which increased both growth rates and per capita fecundity. Size-dependent intercohort competition may have strong impacts on cannibal-victim in- teractions when victims share resources with cannibals. Furthermore, habitat heterogeneity, combined with habitat selection, may limit the extent to which cannibals have a stabilizing effect on population dynamics. Finally, the energy gained by cannibals may have important consequences on population dynamics as this energy is allocated into new recruits.

Gigantic cannibals driving a whole-lake trophic cascade

Trophic cascades have been a central paradigm in explaining the structure of ecological communities but have been demonstrated mainly through comparative studies or experimental manipulations. In contrast, evidence for shifts in trophic cascades caused by intrinsically driven population dynamics is meager. By using empirical data of a cannibalistic fish population covering a 10-year period and a size-structured population model, we show the occurrence of a dynamic trophic cascade in a lake ecosystem, in which the community over time alternates between two different configurations. The intrinsically driven change in the size structure of the fish population from a dominance of stunted individuals to a dominance of gigantic cannibals among adult individuals is the driving force behind distinct abundance switches observed in zooplankton and phytoplankton. The presence of the phase with gigantic cannibals depends critically on the energy they extract from their victims, allowing strong reproduction for a number of years.

Size-dependent foraging efficiency, cannibalism and zooplankton community structure

Abstract To examine size-dependent food web interactions in systems with cannibalism, we compared the abundances of zooplankton and phytoplankton over 2 years in four lakes with cannibalistic perch (Perca fluviatilis) of which two also supported the top predator pike (Esox lucius). The abundance of perch 2 years and older was lower in lakes with pike than in lakes with only perch. In contrast, the abundance of small perch (young-of-the-year and 1-year old) was lower in lakes with only perch suggesting that intense cannibalism reduced these size classes to low levels in lakes lacking pike. Functional response experiments with differently sized perch and zooplankton showed that the attack rate of small perch susceptible to cannibalism was much higher than that of large cannibalising perch. The optimal body size of perch with respect to attack rate was also lower for small zooplankton prey than for large zooplankton. The zooplankton communities in lakes with only perch were dominated by the relatively small species Ceriodaphnia quadrangula and Bosmina spp. and total zooplankton biomass was higher in these lakes than in lakes with both pike and perch. In contrast, the mean size of cladoceran zooplankton was largest in lakes with both pike and perch owing to a dominance of the large zooplankton species Holopedium gibberum in these lakes. We relate these patterns to (1) the low foraging efficiency of large perch on small zooplankton and (2) the low abundance of small zooplanktivorous perch (due to cannibalism) in lakes with only perch. The differences in zooplankton community structure also resulted in different seasonal dynamics of phytoplankton between lakes. Cannibalism introduces a vertical heterogeneity to food webs that causes consumer-resource dynamics that are not predictable from linear food chain models.

Resource limitation during early ontogeny: constraints induced by growth capacity in larval and juvenile fish

Abstract The presence of and mechanisms behind density-dependent growth and resource limitation in larval and juvenile stages of organisms with high mortality such as fish are much debated. We compare observed consumption and growth rates with maximum consumption and growth rates to study the extent of resource limitation in young-of-the-year (YOY) roach (Rutilus rutilus) and perch (Perca fluviatilis). Diet, habitat use, consumption rate and growth rate were measured under varying YOY fish densities over 2 years in four lakes. In the first year, YOY roach and perch were studied under allopatric conditions. Experimental addition of perch roe in the second year also allowed study of YOY of the two species under sympatric conditions in two of the lakes. The diet of YOY roach was dominated by cladoceran zooplankton and YOY roach habitat use was restricted to the shore region in both years. This restricted habitat use did not involve any cost in foraging gain in the first year as consumption and growth rates were very close to maximum rates. During the second year, when the two species coexisted, resources were limited in late season, more so in the littoral than in the pelagic habitat in one lake while the reverse was the case in the other lake. The diet of YOY perch was also dominated by zooplankton, and with increasing perch size the proportion of macroinvertebrate prey in the diet increased. After hatching, YOY perch first utilized the pelagic habitat restricting their habitat use to the shore after 1 to several weeks in the pelagic zone. During the larval period, perch were not resource limited whereas juvenile perch were resource limited in both years. The fact that YOY perch were more resource limited than YOY roach was related to the higher handling capacity and lower attack rate of perch relative to roach, rendering perch more prone to resource limitation. Estimates of resource limitation based on consumption rates and growth rates yielded similar results. This supports the adequacy of our approach to measure resource limitation and suggests that this method is useful for studying resource limitation in organisms with indeterminate growth. Our results support the view that density-dependent growth is rare in larval stages. We suggest that density-dependent growth was absent because larval perch and roach were feeding at maximum levels over a wide range of larvae densities.

Interactions among size-structured populations in a whole-lake experiment: size- and scale-dependent processes

Interactions in size-structured populations are characterized by a mixture of predatory and competitive interactions dependent on the size of the individual organism. We analyzed this mixture of size-structured interactions for a species constellation consisting of perch (Perca fluviatilis) and roach (Rutilus rutilus) in a replicated whole lake experiment over 4 yr. Roach are preyed upon by large perch, but at the same time compete with small perch for zooplankton. Predictions regarding the effects of roach on perch performance and resource dynamics in the whole-lake experiment were based on results from previous pond and enclosure experiments carried out over short (months) time periods. Variables measured in the whole-lake experiment included both individual level parameters (diet and growth of perch) as well as population level parameters (mortality, population numbers and size structures of perch, abundance and biomass of zooplankton and benthic macroinvertebrates). The population size of perch ≥ 2 yr old decreased to less than 10% of the pre-treatment levels in treatment lakes after the introduction of roach and remained low over the whole study period. The mortality in perch one-year-old and older increased with size. Larger perch had a higher growth decrease than smaller perch, but also the individual growth rate of young-of-the-year (YOY) perch was lower in treatment lakes than in control lakes. YOY perch were almost excluded during the winter and spring following the roach introduction. The few perch of this year class that survived had the following years a higher growth than the corresponding year class in control lakes. This could be related to an increased availability of predator-sensitive macroinvertebrates in treatment lakes. Perch in treatment lakes fed to a larger extent on macroinvertebrates and less on pelagic zooplankton than perch in control lakes. As expected, zooplankton abundances decreased in treatment lakes the year following the roach introduction, but thereafter zooplankton abundances were higher in treatment lakes due to the absence of perch recruitment in these lakes. No incidence of perch predation on roach was recorded during the study period. We attribute this to the spatial scale dependent escape ability and habitat use of vulnerable size classes of roach. YOY perch had a stronger impact on between-year variation in zooplankton than YOY roach and this can be related to the higher foraging (handling) capacity of the former and the restricted habitat use of the latter. Individual level parameters (diet, growth) had a higher statistical power than population level parameters (abundance, biomass, mortality), and small-scale experiments successfully predicted a higher proportion (81%) of the individual level parameters. However, the lower success in predicting population level responses (38%) was directly connected to a failure to predict individual level parameters (large perch diet and growth). Our results suggest that the small spatial scale was the major limitation on the predictive ability of the small-scale experiments.