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2.
  • De Roos, André M, et al. (författare)
  • Simplifying a physiologically structured population model to a stage-structured biomass model.
  • 2008
  • Ingår i: Theoretical Population Biology. - : Elsevier Inc.. - 0040-5809 .- 1096-0325. ; 73:1, s. 47-62
  • Tidskriftsartikel (refereegranskat)abstract
    • 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.
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3.
  • Andersson, Jens, et al. (författare)
  • Plastic reources polymorphism : effects or resource availability on Arctic char (Salvelinus alpinus) morphology
  • 2005
  • Ingår i: Biological Journal of the Linnean Society. - London : Acad. P.. - 0024-4066 .- 1095-8312. ; 85:3, s. 341-351
  • Tidskriftsartikel (refereegranskat)abstract
    • Resource polymorphism has been suggested to be a platform for speciation. In some cases resource polymorphism depends on phenotypic plasticity but in other cases on genetic differences between morphotypes, which in turn has been suggested to be the ongoing development of a species pair. Here we study environmentally induced morphological differences in two age classes of Arctic char (Salvelinus alpinus) influencing char performance and diet in relation to resource availability. We found that structurally complex habitats with relatively lower zooplankton densities gave rise to individuals with a deeper body, and a downward positioned tip of the snout compared with individuals from structurally simple habitats with relatively higher zooplankton densities for both age classes. Environment also had an effect on foraging efficiency on zooplankton, with fish from structurally simple habitats had a higher foraging rate than fish from structurally complex habitats. Diet analyses showed that resource use in char mainly depends on the relative abundance of different resources. Therefore, to gain further understanding of resource polymorphism we suggest that future studies must include population dynamic feedbacks by the resources on the consumers.
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5.
  • Andersson, Jens, 1965- (författare)
  • The development of resource polymorphism – Effects of diet, predation risk and population dynamical feedbacks.
  • 2005
  • Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
    • This thesis deals with the evolution of individuals within a species adapted to utilize specific resources, i.e. resource polymorphism. Although a well-known phenomenon, the understanding of the mechanisms behind is not complete. Considering the ruling theories, resource polymorphism is suggested to depend on severe competition for resources, the presence of open niches to be occupied leading to a reduction in competition, and disruptive selection where generalist are out-competed due trade-offs in foraging efficiency for different prey. In order to study resource polymorphism, I have used fish as the animal group in focus and the methods I have used range over laboratory experiments, field experiments, literature surveys and theoretical modelling. In my work, I have showed that different resource use induces different body shapes and that the rate of change is dependent of the encounter rate of different resources. The induced body changes partly led to increased foraging efficiency but surprisingly I did not find any trade-offs due to specialization. However, when studying predation risk in relation to resource polymorphism, my studies point towards that resource use and predation risk may act as balancing factors in such a way that disruptive selection can take place. My work also shows that population feedbacks have to be explored when considering the evolution of resource polymorphism. In pond and field experiments, I found that changes in resource densities affected the actual resource use despite previous adaptations to certain resources. By performing a literature survey, I found that cannibalism indirectly by its effect on population dynamics seems to facilitate the evolution of resource polymorphism. Modelling a size-structured population, I found that resource dynamics were stabilized, and the relative availability of different resources was levelled out due to cannibalism. Taken together, my studies strongly suggest that to understand the development of resource polymorphism in consumer populations, future studies have to include the effect of a dynamic environment both with respect to resources and predators.
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8.
  • Claessen, David, et al. (författare)
  • The effect of population size and recombination on delayed evolution of polymorphism and speciation in sexual populations
  • 2008
  • Ingår i: American Naturalist. - : University of Chicago Press. - 0003-0147 .- 1537-5323. ; 172:1, s. E18-34
  • Tidskriftsartikel (refereegranskat)abstract
    • Recent theory suggests that absolute population size may qualitatively influence the outcome of evolution under disruptive selection in asexual populations. Large populations are predicted to undergo rapid evolutionary branching; however, in small populations, the waiting time to branching increases steeply with decreasing abundance, and below a critical size, the population remains monomorphic indefinitely. Here, we (1) extend the theory to sexual populations and (2) confront its predictions with empirical data, testing statistically whether lake size affects the level of resource polymorphism in arctic char (Salvelinus alpinus) in 22 lakes of different sizes. For a given level of recombination, our model predicts qualitatively similar relations between population size and time to evolutionary branching (either speciation or evolution of genetic polymorphism) as the asexual model, while recombination further increases the delay to branching. The loss of polymorphism at certain loci, an inherent aspect of multilocus-trait evolution, may increase the delay to speciation, resulting in stable genetic polymorphism without speciation. The empirical analysis demonstrates that the occurrence of resource polymorphism depends on both lake size and the number of coexisting fish species. For a given number of coexisting species, the level of polymorphism increases significantly with lake size, thus confirming our model prediction.
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  • De Roos, Andre M., et al. (författare)
  • Ontogenetic symmetry and asymmetry in energetics
  • 2013
  • Ingår i: Journal of Mathematical Biology. - : Springer Science and Business Media LLC. - 0303-6812 .- 1432-1416. ; 66:4-5, s. 889-914
  • Tidskriftsartikel (refereegranskat)abstract
    • Body size ( biomass) is the dominant determinant of population dynamical processes such as giving birth or dying in almost all species, with often drastically different behaviour occurring in different parts of the growth trajectory, while the latter is largely determined by food availability at the different life stages. This leads to the question under what conditions unstructured population models, formulated in terms of total population biomass, still do a fair job. To contribute to answering this question we first analyze the conditions under which a size-structured model collapses to a dynamically equivalent unstructured one in terms of total biomass. The only biologically meaningful case where this occurs is when body size does not affect any of the population dynamic processes, this is the case if and only if the mass-specific ingestion rate, the mass-specific biomass production and the mortality rate of the individuals are independent of size, a condition to which we refer as "ontogenetic symmetry". Intriguingly, under ontogenetic symmetry the equilibrium biomass-body size spectrum is proportional to 1/size, a form that has been conjectured for marine size spectra and subsequently has been used as prior assumption in theoretical papers dealing with the latter. As a next step we consider an archetypical class of models in which reproduction takes over from growth upon reaching an adult body size, in order to determine how quickly discrepancies from ontogenetic symmetry lead to relevant novel population dynamical phenomena. The phenomena considered are biomass overcompensation, when additional imposed mortality leads, rather unexpectedly, to an increase in the equilibrium biomass of either the juveniles or the adults (a phenomenon with potentially big consequences for predators of the species), and the occurrence of two types of size-structure driven oscillations, juvenile-driven cycles with separated extended cohorts, and adult-driven cycles in which periodically a front of relatively steeply decreasing frequencies moves up the size distribution. A small discrepancy from symmetry can already lead to biomass overcompensation; size-structure driven cycles only occur for somewhat larger discrepancies.
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11.
  • de Roos, André M., et al. (författare)
  • Stage-specific predator species help each other to persist while competing for a single prey
  • 2008
  • Ingår i: Proceedings from the National Academy of Science of the United States of America. - Washington, USA : The National Academy of Sciences of the United States of America. - 0027-8424 .- 1091-6490. ; 105:37, s. 13930-13935
  • Tidskriftsartikel (refereegranskat)abstract
    • 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 even not be able to persist at all unless its competitor is also present. The competitor thus increases significantly 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.
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12.
  • Gårdmark, Anna, et al. (författare)
  • Regime shifts in exploited marine food webs : detecting mechanisms underlying alternative stable states using size-structured community dynamics theory
  • 2015
  • Ingår i: Philosophical Transactions of the Royal Society of London. Biological Sciences. - : The Royal Society. - 0962-8436 .- 1471-2970. ; 370:1659, s. 20130262-
  • Tidskriftsartikel (refereegranskat)abstract
    • Many marine ecosystems have undergone 'regime shifts', i.e. abrupt reorganizations across trophic levels. Establishing whether these constitute shifts between alternative stable states is of key importance for the prospects of ecosystem recovery and for management. We show how mechanisms underlying alternative stable states caused by predator-prey interactions can be revealed in field data, using analyses guided by theory on size-structured community dynamics. This is done by combining data on individual performance (such as growth and fecundity) with information on population size and prey availability. We use Atlantic cod (Gadus morhua) and their prey in the Baltic Sea as an example to discuss and distinguish two types of mechanisms, 'cultivation-depensation' and 'overcompensation', that can cause alternative stable states preventing the recovery of overexploited piscivorous fish populations. Importantly, the type of mechanism can be inferred already from changes in the predators' body growth in different life stages. Our approach can thus be readily applied to monitored stocks of piscivorous fish species, for which this information often can be assembled. Using this tool can help resolve the causes of catastrophic collapses in marine predatory-prey systems and guide fisheries managers on how to successfully restore collapsed piscivorous fish stocks.
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13.
  • Hin, Vincent, et al. (författare)
  • Coexistence of Predator and Prey in Intraguild Predation Systems with Ontogenetic Niche Shifts
  • 2011
  • Ingår i: American Naturalist. - Chicago : University of Chicago Press. - 0003-0147 .- 1537-5323. ; 178:6, s. 701-714
  • Tidskriftsartikel (refereegranskat)abstract
    • 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.
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14.
  • Jansson, Mats, et al. (författare)
  • Terrestrial carbon and intraspecific size-variation shape lake ecosystems
  • 2007
  • Ingår i: Trends in Ecology & Evolution. - : Elsevier BV. - 0169-5347 .- 1872-8383. ; 22:6, s. 316-322
  • Forskningsöversikt (refereegranskat)abstract
    • Conceptual models of lake ecosystem structure and function have generally assumed that energy in pelagic systems is derived from in situ photosynthesis and that its use by higher trophic levels depends on the average properties of individuals in consumer populations. These views are challenged by evidence that allochthonous subsidies of organic carbon greatly influence energy mobilization and transfer and the trophic structure of pelagic food webs, and that size variation within consumer species has major ramifications for lake community dynamics and structure. These discoveries represent conceptual shifts that have yet to be integrated into current views on lake ecosystems. Here, we assess key aspects of energy mobilization and size-structured community dynamics, and show how these processes are intertwined in pelagic food webs.
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  • Persson, Lennart, et al. (författare)
  • Cannibalism in a size-structured population : energy extraction and control
  • 2004
  • Ingår i: Ecological Monographs. - : Wiley. - 0012-9615 .- 1557-7015. ; 74:1, s. 135-157
  • Tidskriftsartikel (refereegranskat)abstract
    • Recent size-structured cannibalistic models point to the importance of the energy gain by cannibals and also show that this gain may result in the emergence of giant individuals. We use a combination of a 10-year field study of a perch (Perca fluviatilis) population and quantitative within-season modeling of individual and population-level dynamics to investigate which mechanisms are most likely to drive the dynamics of the studied perch population. We focused on three main aspects to explain observed discrepancies between earlier model predictions and data: (1) introduction of more than one shared resource between cannibals and victims, (2) whether or not several victim age cohorts are necessary to allow giant growth, and (3) the intensity of inter-cohort competition between young-of-the-year (YOY) perch and 1-yr-old perch. At the start of the study period, the perch population was dominated by “stunted” perch individuals, and recruitment of perch to an age of 1-yr-old was negligible. Following a major death in adult perch, strong recruitments of perch to 1-yr-old were thereafter observed for a number of years. As 1-yr-olds these successful recruiters subsequently starved to death due to competition with the new YOY. The few surviving adult perch accelerated substantially in growth and became “giants.” At the end of the study period, the perch population moved back to the situation with stunted individuals. There was a high agreement between observed diets of cannibalistic perch and those predicted by the model for both the stunted and the giant phases. Analyses of growth rates showed that cannibalistic perch could become giants on a diet of YOY perch only, but that a supplement with the second shared resource (macroinvertebrates) was needed to reach the observed sizes. Modeling of growth and diet in the giant phase showed an exploitative competitive effect of YOY perch on 1-yr-old perch, but a restriction in habitat use of 1-yr-old perch had to be assumed to yield the observed growth rate and diet. The resource dynamics of zooplankton and macroinvertebrates were both accurately predicted by the model. Also, YOY perch mortality was accurately predicted and, furthermore, suggested that one of the trawling methods used may underestimate the number of YOY perch when they increase in size. We conclude that the presence of a second shared resource and the restricted habitat use and absence of cannibalistic consumption by 1-yr-old perch individuals are two important mechanisms to explain the discrepancy between model predictions and data. Our results also point to the fact that that the dynamics observed may be explained by complex dynamics not involving the presence of a giant and dwarf cycle.
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  • Persson, Lennart, et al. (författare)
  • Density-dependent interactions in an Arctic char - brown trout system : competition, predation, or both?
  • 2013
  • Ingår i: Canadian Journal of Fisheries and Aquatic Sciences. - : Canadian Science Publishing. - 0706-652X .- 1205-7533. ; 70:4, s. 610-616
  • Tidskriftsartikel (refereegranskat)abstract
    • In the study of mechanisms structuring fish communities, mixed competition-predation interactions where large predators feed on prey fish versus those in which small predators compete with prey fish for a shared prey have been the focus of substantial research. We used a long-term data set from a system inhabited by brown trout (Salmo trutta) (predator) and Arctic char (Salvelinus alpinus) (prey) to evaluate whether mixed interspecific interactions were present in this system as suggested in other studies focusing on this species pair. We found no evidence for a negative interspecific density dependence in individual performance in either Arctic char or brown trout. In contrast, a negative intraspecific density dependence was present, especially in Arctic char. Furthermore, large brown trout condition showed a positive response to encounter rate with Arctic char (related to the density of small Arctic char). The most parsimonious interaction module to explain the Arctic char - brown trout interaction patterns in the studied system does therefore not need to include interspecific competition. We suggest that size-structured mixed competition-predation interactions in different systems are realized as being either mainly structured through interspecific predation or by competition depending on species life history characteristics and environmental conditions.
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19.
  • Persson, Lennart, et al. (författare)
  • Food-dependent individual growth and population dynamics in fishes
  • 2006
  • Ingår i: Journal of fish biology. - Oxford : Blackwell Publishing. - 0022-1112 .- 1095-8649. ; , s. 1-20
  • Konferensbidrag (refereegranskat)abstract
    • It is long since well established that growth and development in fish individuals are heavily dependent on food intake. Yet, this dependence of individual development on food levels has only to a limited extent been taken into consideration when Studying fish population and community processes. Using the modelling framework of physiologically Structured population models and empirical data For a number of species configurations, how different size-dependent processes may affect fish population dynamics and community structures are reviewed. Considering competitive interactions, cohort interactions will often give rise to cohort cycles driven by an inequality in competitive abilities between differently sized individuals. The addition of cannibalism may dampen these cycles, the extent to which is dependent on life-history characteristics of the cannibals. The circumstance that individuals Change their trophic position over their life cycle as a result of an increase in size gives rise to life history omnivory. In such omnivorous systems, food-dependent growth demotes the potential for predatory and prey fishes to coexist. In tritrophic food chains, food-dependent growth in the intermediate consumer may lead to the presence of bistability including sensitivity to catastrophic behaviour. These results shed new light on the drastic decreases observed in the stocks of many marine fish top predators including their inability to recover after fishing moratoria, and on the suggested presence of alternative states in freshwater fish communities.
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20.
  • Persson, Lennart, et al. (författare)
  • Mixed competition-predation : potential vs. realized interactions
  • 2012
  • Ingår i: Journal of Animal Ecology. - Malden, MA : Wiley-Blackwell. - 0021-8790 .- 1365-2656. ; 81:2, s. 483-493
  • Tidskriftsartikel (refereegranskat)abstract
    • 1. Life-history omnivory or size-induced mixed competitionpredation systems have under many conditions theoretically been shown to be fragile, whereas at the same time existing empirical data suggest such systems to be common in nature. 2. In a whole lake experiment covering 17 years, we analysed the effects of the introduction of the intraguild prey roach (Rutilus rutilus) on the population size and individual performance of the intraguild predator perch (Perca fluviatilis) and on resource levels in two low productivity systems. 3. A strong long-term effect of roach on the zooplankton resource but not on the macroinvertebrate resource was present. Competitive effects of roach on perch were observed in one of the lakes the first years after the introduction, but at the end of the study no competitive effect of roach on either size class of perch was observed in any of the two lakes. In contrast, a positive predatory effect reflected in improved growth rates of older perch was present. 4. The lack of a support for a competitive effect of roach on small perch raises the question of the importance of mixed competition-predation interactions in life-history omnivorous systems and the problem of comparing descriptive data on feeding relationships with theoretical predictions based on interaction modules.
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  • Persson, Lennart, et al. (författare)
  • State-dependent invasion windows for prey in size-structured predator–prey systems: whole lake experiments
  • 2007
  • Ingår i: Journal of Animal Ecology. - : Wiley. - 0021-8790 .- 1365-2656. ; 76:1, s. 94-104
  • Tidskriftsartikel (refereegranskat)abstract
    • 1.In size-structured communities where individuals grow in size over their life cycle, interactions between species will shift between competitive and predatory interactions depending on size relationships. The outcome of interactions will subsequently depend on the strength of competitive and predatory interactions, respectively.2.In a whole lake experiment including four experimental lakes, it was tested under which conditions the competing prey, roach Rutilus rutilus, could successfully recruit into systems previously occupied by the predator, perch Perca fluviatilis. Two replicated introduction experiments were carried out 3 years apart.3.Roach were able to successfully recruit into three of the four experimental lakes of which two were also inhabited by the top predator pike Esox lucius. Resource levels were unrelated to whether roach could successfully recruit into the systems as recruiting roach in all years were feeding close to their maximum rate.4.High population fecundity of roach and low predation pressure by perch combined were necessary ingredients for successful recruitment and the presence of only one of these conditions did not result in successful recruitment.5.It is hypothesized that, although roach were able to successfully recruit into one lake with only perch present in addition to the two lakes that also inhabited pike, long-term coexistence of roach and perch depends on the presence of another top predator (e.g. pike) selectively preying on perch. This hypothesis was supported by data on co-occurrence of perch and roach in different lakes.6.Overall, the results are in accordance with expectation of size-structured life-history omnivory theory suggesting that coexistence between top predator and intermediate consumer is fragile.
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23.
  • Persson, Lennart, et al. (författare)
  • Symmetry breaking in ecological systems through different energy efficiencies of juveniles and adults
  • 2013
  • Ingår i: Ecology. - : Wiley. - 0012-9658 .- 1939-9170. ; 94:7, s. 1487-1498
  • Tidskriftsartikel (refereegranskat)abstract
    • Ontogenetic development is a fundamental aspect of the life history of all organisms and has major effects on population and community dynamics. We postulate a general conceptual framework for understanding these effects and claim that two potential energetics bottlenecks at the level of the individual organismthe rate by which it develops and the rate by which it reproducesform a fundamental route to symmetry-breaking in ecological systems, leading to ontogenetic asymmetry in energetics. Unstructured ecological theory, which ignores ontogenetic development, corresponds to a limiting case only, in which mass-specific rates of biomass production through somatic growth and reproduction, and biomass loss through mortality, are independent of body size (ontogenetic symmetry). Ontogenetic symmetry results in development and reproduction being limited to the same extent by food density. In all other cases, symmetry-breaking occurs. Ontogenetic asymmetry results in increases in juvenile, adult, or even total biomass in response to mortality. At the community level, this gives rise to alternative stable states via predator-induced shifts in prey size distributions. Ontogenetic asymmetry furthermore leads to two distinct types of cycles in population dynamics, depending on whether development or reproduction is most energy limited. We discuss the mechanisms giving rise to these phenomena and the empirical support for them. We conclude that the concepts of ontogenetic symmetry and ontogenetic asymmetry form a novel and general organizing principle on which future ecological theory should be developed.
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24.
  • Persson, Lennart, et al. (författare)
  • The ecological foundation for ecosystem-based management of fisheries : mechanistic linkages between the individual-, population-, and community-level dynamics
  • 2014
  • Ingår i: ICES Journal of Marine Science. - : Oxford University Press. - 1054-3139 .- 1095-9289. ; 71:8, s. 2268-2280
  • Tidskriftsartikel (refereegranskat)abstract
    • Food-dependent growth and size-dependent interactions form cornerstones in the dynamics of fish populations. Using two freshwater examples, we illustrate the importance of considering both these cornerstones for understanding system dynamics. Moreover, a proper understanding of the dynamics requires mechanistic linkages between individual-, population-, and community-level processes based on mass conservation principles. In one example, we further find that quantitative predictions of individual-level energy flows are essential for understanding the community dynamics. This mechanistic approach to understanding system dynamics is generally not reflected in fisheries models as an overview shows that only half of them incorporate food-dependent growth, and none fully observe the principles of mass conservation. As a marine example we examine patterns in the Baltic Sea system and show that no relationship between cod growth and sprat biomass is present related to the low size resolution in prey fish. Linking individual cod performance to its resource base is complicated by the many prey types cod uses over its life cycle. We conclude that an ecological perspective including size-and food-dependent processes is vital for ecosystem-based fisheries management making necessary a proper description of the interactive trophic structure as a result of mechanistic linkages between individual, population, and community processes.
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25.
  • Reichstein, Birte, 1983-, et al. (författare)
  • Coexistence in a size-structured intraguild predation system promoted by an ontogenetic diet shift in the consumer
  • Annan publikation (övrigt vetenskapligt/konstnärligt)abstract
    • 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. 
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26.
  • Reichstein, Birte, et al. (författare)
  • Exclusive juvenile predator resource promotes coexistence in a size-structured intraguild predation system
  • Annan publikation (övrigt vetenskapligt/konstnärligt)abstract
    • 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. 
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27.
  • Reichstein, Birte, 1983-, et al. (författare)
  • Ontogenetic asymmetry modulates population biomass production and response to harvest
  • 2015
  • Ingår i: Nature Communications. - : Nature Publishing Group. - 2041-1723. ; 6
  • Tidskriftsartikel (refereegranskat)abstract
    • 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.
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28.
  • Reichstein, Birte, 1983- (författare)
  • Ontogenetic bottlenecks : effects on intraguild predation systems and ecosystem efficiency
  • 2015
  • Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
    • 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.
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29.
  • Reichstein, Birte, 1983-, et al. (författare)
  • Predator life history affects persistence times of predators and consumers in an intraguild predation system
  • Annan publikation (övrigt vetenskapligt/konstnärligt)abstract
    • 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.
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30.
  • Schröder, Arne, 1974- (författare)
  • Alternative Stable States in Size-Structured Communities : Patterns, Processes, and Mechanisms
  • 2008
  • Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
    • Alternative stable states have been, based on theoretical findings, predicted to be common in ecological systems. Empirical data from a number of laboratory and natural studies strongly suggest that alternative stable states also occur in real populations, communities and ecosystems. Potential mechanisms involve population size-structure and food-dependent individual development. These features can lead to ontogenetic niche shifts, juvenile recruitment bottlenecks and emergent Allee effects; phenomena that establish destabilising positive feedbacks in a system and hence create alternative stable states.I studied the consequences of population size-structure for community dynamics at different scales of system complexity. I performed laboratory and ecosystem experiments. Small poecilliid fishes and planktonic invertebrates with short generation times and life spans were used as model organisms. This allowed me to assess the long-term dynamics of the populations and communities investigated.The main experimental results are: (a) An ontogenetic niche shift in individuals of the phantom midge Chaoborus made the population vulnerable to an indirect competitive recruitment bottleneck imposed by cladoceran mesozooplankton via rotifers. Consequentially the natural zooplankton food web exhibited two alternative attractors. (b) Body size determined the success of Poecilia reticulata invading resident population of Heterandria formosa and thus the type of alternative stable state that established. Small invaders were outcompeted by the residents, whereas large invaders excluded their competitor by predating on its recruits. (c) External juvenile and adult mortality altered the internal feedback structure that regulates a laboratory population of H. formosa in such a way that juvenile biomass increased with mortality. This biomass overcompensation in a prey population can establish alternative stable states with top-predators being either absent or present.The major conclusion is that size-structure and individual growth can indeed lead to alternative stable states. The considerations of these ubiquitous features of populations offer hence new insights and deeper understanding of community dynamics. Alternative stable states can have tremendous consequences for human societies that utilise the ecological services provided by an ecological system. Understanding the effects of size-structure on alternative stability is thus crucial for sustainable exploitation or production of food resources.
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31.
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32.
  • Schröder, Arne, et al. (författare)
  • Complex shifts between food web states in response to whole-ecosystem manipulations
  • 2012
  • Ingår i: Oikos. - : Wiley. - 0030-1299 .- 1600-0706. ; 121:3, s. 417-427
  • Tidskriftsartikel (refereegranskat)abstract
    • Food webs can respond in surprising and complex ways to temporary alterations in their species composition. When such a perturbation is reversed, food webs have been shown to either return to the pre-perturbation community state or remain in the food web configuration that established during the perturbation. Here we report findings from a replicated whole-lake experiment investigating food web responses to a perturbation and its consecutive reversal. We could identify three distinct community states in the food web that corresponded to the periods before, during and after the perturbation. Most importantly, we demonstrate the establishment of a distinct post-perturbation food web configuration that differed from both the pre- and during-perturbation communities in phytoplankton biomass and micro- and mesozooplankton species composition. We suggest that the pre- and post-perturbation food web configurations may represent two alternative stable community states. We provide explanations for how each of the contrasting communities may be maintained through altered species interactions. These findings add to the discussion of how natural food webs react to environmental change and imply that the range of potential ecosystem dynamics in response to perturbations can be wider and more complex than is often recognized.
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33.
  • Schröder, Arne, et al. (författare)
  • Direct experimental evidence for alternative stable states : a review
  • 2005
  • Ingår i: Oikos. - : Wiley. - 0030-1299 .- 1600-0706. ; 110:1, s. 3-19
  • Tidskriftsartikel (refereegranskat)abstract
    • A large number of studies have presented empirical arguments for the existence of alternative stable states (ASS) in a wide range of ecological systems. However, most of these studies have used non-manipulative, indirect methods, which findings remain open for alternative explanations. Here, we review the direct evidence for ASS resulting from manipulation experiments. We distinguish four conclusive experimental approaches which test for predictions made by the hysteresis effect: (1) discontinuity in the response to an environmental driving parameter, (2) lack of recovery potential after a perturbation, (3) divergence due to different initial conditions and (4) random divergence. Based on an extensive literature search we found 35 corresponding experiments. We assessed the ecological stability of the reported contrasting states using the minimum turnover of individuals in terms of life span and classified the studies according to 4 categories: (1) experimental system, (2) habitat type, (3) involved organisms and (4) theoretical framework. 13 experiments have directly demonstrated the existence of alternative stable states while 8 showed the absence of ASS in other cases. 14 experiments did not fulfil the requirements of a conclusive test, mostly because they applied a too short time scale. We found a bias towards laboratory experiments compared to field experiments in demonstrating bistability. There was no clear pattern of the distribution of ASS over categories. The absence of ASS in 38% of the tested systems indicates that ASS are just one possibility of how ecological systems can behave. The relevance of the concept of ASS for natural systems is discussed, in particular under consideration of the observed laboratory bias, perturbation frequency and variable environments. It is argued, that even for a permanently transient system, alternative attractors may still be of relevance.
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34.
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35.
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36.
  • van Kooten, Tobias, et al. (författare)
  • Population dynamical consequences of gregariousness in a size-structured consumer–resource interaction
  • 2007
  • Ingår i: Journal of Theoretical Biology. - : Elsevier BV. - 0022-5193. ; 254:4, s. 763-774
  • Tidskriftsartikel (refereegranskat)abstract
    • 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.
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37.
  • van Kooten, Tobias, et al. (författare)
  • Size at hatching determines population dynamics and response to harvesting in cannibalistic fish
  • 2010
  • Ingår i: Canadian Journal of Fisheries and Aquatic Sciences. - 0706-652X .- 1205-7533. ; 67:2, s. 401-416
  • Tidskriftsartikel (refereegranskat)abstract
    • We hypothesize that size at hatching strongly affects population dynamics of cannibalistic fish species and is a crucial determinant of how populations respond to selective removal of large individuals (harvesting). We use a mechanistic mathematical model to study the relation between hatching size and response to harvesting mortality, using Eurasian perch (Perca fluviatilis) as a model organism. We show how hatching size determines dynamics through its effect on the relative strength of cannibalistic mortality and resource competition as mechanisms of population regulation. In populations with intermediate and large hatching size, cannibalistic mortality is an important determinant of population dynamics. and harvesting destabilizes population dynamics. When hatching size is small, population stability is less sensitive to this type of harvesting. Populations hatching at small size are regulated by competition, and harvesting large individuals affects such populations less. Harvesting can also induce the growth of very large individuals, absent in unharvested populations. Our results show that harvesting in cannibalistic lake fish populations can strongly alter Population dynamics in ways that can only be anticipated on the basis of mechanistic knowledge about how populations are regulated.
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38.
  • van Kooten, Tobias, et al. (författare)
  • Size-Dependent Mortality Induces Life-History Changes Mediated through Population Dynamical Feedbacks
  • 2007
  • Ingår i: American Naturalist. ; 170:2, s. 258-270
  • Tidskriftsartikel (refereegranskat)abstract
    • 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.
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39.
  • van Leeuwen, Anieke, et al. (författare)
  • Predators with multiple ontogenetic niche shifts have limited potential for population growth and top-down control of their prey
  • 2013
  • Ingår i: American Naturalist. - : University of Chicago Press. - 0003-0147 .- 1537-5323. ; 182:1, s. 53-66
  • Tidskriftsartikel (refereegranskat)abstract
    • Catastrophic collapses of top predators have revealed trophic cascades and community structuring by top-down control. When populations fail to recover after a collapse, this may indicate alternative stable states in the system. Overfishing has caused several of the most compelling cases of these dynamics, and in particular Atlantic cod stocks exemplify such lack of recovery. Often, competition between prey species and juvenile predators is hypothesized to explain the lack of recovery of predator populations. The predator is then considered to compete with its prey for one resource when small and to subsequently shift to piscivory. Yet predator life history is often more complex than that, including multiple ontogenetic diet shifts. Here we show that no alternative stable states occur when predators in an intermediate life stage feed on an additional resource (exclusive to the predator) before switching to piscivory, because predation and competition between prey and predator do not simultaneously structure community dynamics. We find top-down control by the predator only when there is no feedback from predator foraging on the additional resource. Otherwise, the predator population dynamics are governed by a bottleneck in individual growth occurring in the intermediate life stage. Therefore, additional resources for predators may be beneficial or detrimental for predator population growth and strongly influence the potential for top-down community control.
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40.
  • Wollrab, Sabine, et al. (författare)
  • Ontogenetic diet shifts promote predator-mediated coexistence
  • 2013
  • Ingår i: Ecology. - : Ecological Society of America. - 0012-9658 .- 1939-9170. ; 94:12, s. 2886-2897
  • Tidskriftsartikel (refereegranskat)abstract
    • It is widely believed that predation moderates interspecific competition and promotes prey diversity. Still, in models of two prey sharing a resource and a predator, predator-mediated coexistence occurs only over narrow ranges of resource productivity. These models have so far ignored the widespread feature of ontogenetic diet shifts in predators. Here, we theoretically explore the consequences of a diet shift from juvenile to adult predator stages for coexistence of two competing prey. We find that only very minor deviations from perfectly identical diets in juveniles and adults destroy the traditional mechanism of predator-mediated coexistence, which requires an intrinsic trade-off between prey defendedness and competitive ability. Instead, predator population structure can create an emergent competition-predation trade-off between prey, where a bottleneck in one predator stage enhances predation on the superior competitor and relaxes predation on the inferior competitor, irrespective of the latter's intrinsic defendedness. Pronounced diet shifts therefore greatly enlarge the range of prey coexistence along a resource gradient. With diet shifts, however, coexistence usually occurs as one of two alternative states and, once lost, may not be easily restored.
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41.
  • Wollrab, Sabine, et al. (författare)
  • Simple rules describe bottom-up and top-down control in food webs with alternative energy pathways
  • 2012
  • Ingår i: Ecology Letters. - : Wiley-Blackwell. - 1461-023X .- 1461-0248. ; 15:9, s. 935-946
  • Tidskriftsartikel (refereegranskat)abstract
    • Many human influences on the world's ecosystems have their largest direct impacts at either the top or the bottom of the food web. To predict their ecosystem-wide consequences we must understand how these impacts propagate. A long-standing, but so far elusive, problem in this endeavour is how to reduce food web complexity to a mathematically tractable, but empirically relevant system. Simplification to main energy channels linking primary producers to top consumers has been recently advocated. Following this approach, we propose a general framework for the analysis of bottom-up and top-down forcing of ecosystems by reducing food webs to two energy pathways originating from a limiting resource shared by competing guilds of primary producers (e.g. edible vs. defended plants). Exploring dynamical models of such webs we find that their equilibrium responses to nutrient enrichment and top consumer harvesting are determined by only two easily measurable topological properties: the lengths of the component food chains (oddodd, oddeven, or eveneven) and presence vs. absence of a generalist top consumer reconnecting the two pathways (yielding looped vs. branched webs). Many results generalise to other looped or branched web structures and the model can be easily adapted to include a detrital pathway.
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