Detecting alternative attractors in ecosystem dynamics

Saterberg and McCann introduce a time-series method to test whether systems exhibit alternative dynamical attractors. Through simulated and experimental data from a planktonic predator-prey system, their results show that if population dynamics are induced by internal factors to the system, then alternative dynamical attractors can be detected.Dynamical systems theory suggests that ecosystems may exhibit alternative dynamical attractors. Such alternative attractors, as for example equilibria and cycles, have been found in the dynamics of experimental systems. Yet, for natural systems, where multiple biotic and abiotic factors simultaneously affect population dynamics, it is more challenging to distinguish alternative dynamical behaviors. Although recent research exemplifies that some natural systems can exhibit alternative states, a robust methodology for testing whether these constitute distinct dynamical attractors is currently lacking. Here, using attractor reconstruction techniques we develop such a test. Applications of the methodology to simulated, experimental and natural time series data, reveal that alternative dynamical behaviors are hard to distinguish if population dynamics are governed by purely stochastic processes. However, if population dynamics are brought about also by mechanisms internal to the system, alternative attractors can readily be detected. Since many natural populations display evidence of such internally driven dynamics, our approach offers a method for empirically testing whether ecosystems exhibit alternative dynamical attractors

Ecological impact of changes in intrinsic growth rates of species at different trophic levels

Decreased and increased intrinsic growth rate and abundance of a single species can severely and negatively impact other species in the same food web. Here we compare the wider system effects of decreased and increased intrinsic growth rates of species occupying different trophic levels. Specifically, we derive the change in growth rate of a single (focal) species necessary to cause a 90% reduction in the abundance - a quasi-extinction - of another species in model communities. We find that even relatively small changes, negative as well as positive, in the growth rate of the focal species can cause quasi-extinctions of others. Furthermore, the magnitude of change needed to cause a quasi-extinction depends on the trophic level of the perturbed species. The potential ecosystem impact of such 'negative' and 'positive' changes is largely unknown. We argue that such a targeted decrease or increase could be induced by human interference, such as hunting or harvesting, but also by an outbreak or fade-out of an infectious disease. As ecosystems maintain many and diverse infectious agents, these results suggest that these agents may play an important role in the structure and balance of ecosystems

Species- and origin-specific susceptibility to bird predation among juvenile salmonids

Juvenile salmonids often experience high mortality rates during migration and bird predation is a common source of mortality. Research suggests that hatchery-reared salmonids are more prone to predation than wild salmonids, and that Atlantic salmon (Salmo salar) experience lower predation than Sea trout (Salmo trutta), yet telemetry studies have displayed equivocal results. Here, using a large data set on passive integrated transponder (PIT) tagged hatchery-reared and wild juveniles of Atlantic salmon and Sea trout (25,769 individuals) we investigate predation probability by piscivorous birds (mainly Great Cormorants Phalarocorax carbo) on salmonids originating from River Dalälven in Sweden. Bird colonies and roosting sites were scanned annually (2019–2021), and the temporal dynamics of bird predation on salmonids released in 2017–2021 was assessed. Hatchery-reared trout was clearly most susceptible to cormorant predation (0.31, 90% credibility interval [CRI] = 0.14–0.53), followed by wild trout (0.19, 90% CRI = 0.08–0.37), hatchery-reared salmon (0.13, 90% CRI = 0.07–0.23), and wild salmon (0.08, 90% CRI = 0.04–0.14), in subsequent order. This order in predation probability was consistent across all studied tag- and release-years, suggesting that the opportunistic foraging of cormorants affects the overall survival of juvenile salmonids, but that the inherent predation risk between different salmonid types differs systematically

Ecologically Sustainable Exploitation Rates - a multispecies approach for fisheries management

Fisheries management is slowly evolving from its traditional single-species focus to a more holistic ecosystem-based approach. Yet, limits for exploitation are almost always set based on single-species models, treating species as isolated entities. This is problematic since the sustainability of a fishery hinges on its effects on the exploited community as a whole. Here, we develop a novel analytical approach of estimating exploitation rates that are sustainable with respect to the state of whole fish communities. Our approach simultaneously addresses species interactions, environmental covariates and natural variability of population sizes, yet it is framed around a simple and accessible objective. We derive Ecologically Sustainable Exploitation Rates, that is exploitation rates associated with a maximum acceptable probability (determined by management) that any interacting species decreases to an unacceptably low population size. Using models fitted to an exploited fish community, we show how accounting for species interactions constrains the possibilities for ecologically sustainable exploitation. The conventional omission of species interactions may thus result in overestimated exploitation limits. Moreover, our application rendered a counterintuitive result: it suggests that the exploitation of one species should increase, as compared to mean historical levels, for the purpose of conservation of the community as a whole. Such insights could impossibly be gained using single-species approaches, illustrating the need to adopt multispecies models in fisheries management. Analytical derivation of Ecologically Sustainable Exploitation Rates offers a mean to do so

Functional Extinctions of Species in Ecological Networks

Current rates of extinctions are estimated to be around 1000 times higher than background rates that would occur without anthropogenic impacts. These extinction rates refer to the traditional view of extinctions, i.e. numerical extinctions. This thesis is about another type of extinctions: functional extinctions. Those occur when the abundance of a species is too small to uphold the species’ ecologically interactive role. I have taken a theoretical approach and used dynamical models to investigate functional extinctions and threshold values for species’ mortality rates in ecological networks. More specifically, I have derived threshold values for focal species mortality rates at which another species or the focal species itself goes numerically extinct (Paper I-II), or transgresses some predefined threshold abundance (Paper III). If an increased mortality rate of a focal species causes another species to go numerically extinct, the focal species can be regarded as functionally extinct, since its abundance is no longer large enough to uphold its ecologically interactive role. Such functional extinctions are investigated in the first papers (Paper I-II). In the following paper, limits for both increased and decreased mortality rates of species are explored (Paper III). Paper III also extends the basic theoretical idea developed in paper I-II into a more applied setting. In this paper I develop a time series approach aimed at estimating fishing mortalities associated with a low risk that any species in a community transgresses some predefined critical abundance threshold. In the last paper (Paper IV) the community wide effect of changes in the abundance of species is investigated. In the first paper (Paper I) I investigate threshold levels for the mortality rate of species in ecological networks. When an increased mortality rate of a focal species causes another species to go extinct, the focal species can be characterized as functional extinct, even though it still exists. Such functional extinctions have been observed in a few systems, but their frequency and general patterns have been unexplored. Using a new analytical method the patterns and frequency of functional extinctions in theoretical and empirical ecological networks are explored. It is found that the species most likely to be the first to go extinct is not the species whose mortality rate is increased, but instead another species in the network. The species which goes extinct is often not even directly linked to the species whose mortality rate is increased, but instead indirectly linked. Further, it is found that large-bodied species at the top of food chains can only be exposed to small increases in mortality rate and small decreases in abundance before going functionally extinct compared to small-bodied species lower in the food chains. These results illustrate the potential importance of functional extinctions in ecological networks and lend support to arguments advocating a more community-oriented approach in conservation biology, with target levels for populations based on ecological functionality rather than the mere persistence of species. In Paper II I use the approach developed in Paper I to explore the frequency and patterns of functional extinctions in ecological networks with varying proportions of mutualistic and antagonistic (predator-prey) interactions. The general results from Paper I are also found in Paper II; that is, an increased mortality rate of one focal species often first leads to an extinction of another species rather than to an extinction of the focal species itself. Further, the frequency of functional extinctions is higher in networks containing a mixture of interaction types than in networks with only antagonistic interactions. Overall, this study generalize the findings of paper I for networks containing a variety of interaction types. To make the theoretical approaches developed in paper I-II operational in a management setting I develop a time series approach aimed at estimating ecologically sustainable fishing mortalities in a multispecies fisheries context (Paper III). An ecologically sustainable fishing mortality is here defined as a long-term fishing mortality associated with a multispecies objective which infers a low risk that any species, either the focal species itself or another species, in a community transgresses a critical biomass limit, below which the risk of recruitment failure is high. The approach is exemplified using a statistical food web model of the dominating fish stocks in the Baltic Sea. For the most abundant fish stock a counterintuitive result is found; it is more likely that the multispecies objective is met if its mortality caused by fishing is increased compared to if it is decreased. Further, simultaneous changes of the fishing mortality of a number of interacting species in the food web model shows a much narrower region of possible sustainable fishing mortalities than a single species approach, something that is not captured by current stock assessment models. Altogether these results are governed by indirect effects propagating in the community and pinpoints the need to adopt community dynamical approaches in fisheries management. The population sizes of many species in the world are declining. Negative population trends are particular pronounced in large-bodied herbivores and carnivores, species known to play important regulatory roles in many ecosystems. Although this indicates that the ecological consequence of declining populations of species might be profound, its impact on ecosystem stability remains largely unexplored. In paper IV it is therefore explored how declining populations of rare and common species affects the resilience – recovery rate – of ecological networks. An analytical approximation shows that network resilience is a function of the harmonic mean of the species’ abundances. This means that network resilience is especially sensitive to declining abundances of rare species. Consistent with this analytically derived result, a clear and positive relationship between resilience and the abundance of the rarest species in a broad spectrum of dynamical models of ecological networks is found. Together these results illustrate the potentially negative consequences of declining populations of rare species for the stability of the ecological systems in which they are embedded, and provide ecological arguments for the protection and management of rare species

Minimum Ecologically Viable Populations : Risk assessment from a multispecies perspective

The extinction risk of threatened species has traditionally been assessed by the use of tools of Population Viability Analysis (PVA). Species interactions, however, have seldom been accounted for in PVA:s. The omission of species interactions in risk assessments may further lead to serious mistakes when setting target sizes of populations. Even a slight abundance decrease of a target species may result in changes of the community structure; in the worst case leading to a highly impoverished community. Of critical importance to conservation is therefore the question of how many individuals of a certain population that is needed in order to avoid this kind of consequences. In the current study, a stochastic multispecies model is used to estimate minimum ecological viable populations (MEVP); earlier defined as “the minimum size of a population that can survive before itself or some other species in the community becomes extinct”. The MEVP:s are compared to population sizes given by a single species model where interactions with other species are treated as a constant source incorporated in the species specific growth rate. MEVP:s are found to be larger than the population sizes given by the single species model. The results are trophic level dependent and multispecies approaches are suggested to be of major importance when setting target levels for species at the basal level. Species at higher trophic levels, however, are altogether more prone to extinction than species at the basal level, irrespective of food web size and food web complexity

Minimum Ecologically Viable Populations : Risk assessment from a multispecies perspective

The extinction risk of threatened species has traditionally been assessed by the use of tools of Population Viability Analysis (PVA). Species interactions, however, have seldom been accounted for in PVA:s. The omission of species interactions in risk assessments may further lead to serious mistakes when setting target sizes of populations. Even a slight abundance decrease of a target species may result in changes of the community structure; in the worst case leading to a highly impoverished community. Of critical importance to conservation is therefore the question of how many individuals of a certain population that is needed in order to avoid this kind of consequences. In the current study, a stochastic multispecies model is used to estimate minimum ecological viable populations (MEVP); earlier defined as “the minimum size of a population that can survive before itself or some other species in the community becomes extinct”. The MEVP:s are compared to population sizes given by a single species model where interactions with other species are treated as a constant source incorporated in the species specific growth rate. MEVP:s are found to be larger than the population sizes given by the single species model. The results are trophic level dependent and multispecies approaches are suggested to be of major importance when setting target levels for species at the basal level. Species at higher trophic levels, however, are altogether more prone to extinction than species at the basal level, irrespective of food web size and food web complexity

Ecological impact of changes in intrinsic growth rates of species at different trophic levels

Decreased and increased intrinsic growth rate and abundance of a single species can severely and negatively impact other species in the same food web. Here we compare the wider system effects of decreased and increased intrinsic growth rates of species occupying different trophic levels. Specifically, we derive the change in growth rate of a single (focal) species necessary to cause a 90% reduction in the abundance – a quasi-extinction – of another species in model communities. We find that even relatively small changes, negative as well as positive, in the growth rate of the focal species can cause quasi-extinctions of others. Furthermore, the magnitude of change needed to cause a quasi-extinction depends on the trophic level of the perturbed species. The potential ecosystem impact of such ‘negative' and ‘positive' changes is largely unknown. We argue that such a targeted decrease or increase could be induced by human interference, such as hunting or harvesting, but also by an outbreak or fade-out of an infectious disease. As ecosystems maintain many and diverse infectious agents, these results suggest that these agents may play an important role in the structure and balance of ecosystems

Ecological impact of changes in intrinsic growth rates of species at different trophic levels

Decreased and increased intrinsic growth rate and abundance of a single species can severely and negatively impact other species in the same food web. Here we compare the wider system effects of decreased and increased intrinsic growth rates of species occupying different trophic levels. Specifically, we derive the change in growth rate of a single (focal) species necessary to cause a 90% reduction in the abundance – a quasi-extinction – of another species in model communities. We find that even relatively small changes, negative as well as positive, in the growth rate of the focal species can cause quasi-extinctions of others. Furthermore, the magnitude of change needed to cause a quasi-extinction depends on the trophic level of the perturbed species. The potential ecosystem impact of such ‘negative' and ‘positive' changes is largely unknown. We argue that such a targeted decrease or increase could be induced by human interference, such as hunting or harvesting, but also by an outbreak or fade-out of an infectious disease. As ecosystems maintain many and diverse infectious agents, these results suggest that these agents may play an important role in the structure and balance of ecosystems

The contribution of rare species to a community\u27s resilience

The BES Quantitative Ecology SIG, St Andrews, Scotland, 9 July 2018Theoretical results for generalised Lotke-Volterra systems that demonstrate the role of rare species in determining the characteristic return time of the system.Science Foundation Irelan