Cannibalism facilitated by parasite infection induces dispersal in a semi-aquatic insect.

Parasites are known to have direct effects on host dispersal ability and motivation. Yet, parasites have a variety of impacts on host populations, including shaping predation and cannibalism rates, and therefore may also have indirect effects on host dispersal; these indirect pathways have not been studied. We tested the hypothesis that parasites influence host dispersal through effects on cannibalism using backswimmers (Notonecta undulata) and Hydrachnidia freshwater mites. Mite parasitism impedes swimming in backswimmers, which we found increased their vulnerability to cannibalism. We imposed a manipulation that varied cannibalism rates across experimental populations consisting of a mix of backswimmers with and without simulated parasites. Using simulated parasites allowed us to examine the effects of cannibalism without introducing infection risk. We found that the odds of dispersal for infected backswimmers increased by 2.25× with every 10% increase in the risk of being cannibalized, and the odds of dispersal for healthy backswimmers increased by 2.34× for every additional infected backswimmer they consumed. Our results suggest that cannibalism was used as an energy source for dispersal for healthy individuals, while the risk of being eaten motivated dispersal in infected individuals. These results elucidate the complex ways that parasites impact host populations and strengthen our understanding of host-parasite interactions, including host and parasite population stability and spread. This article is part of the theme issue 'Diversity-dependence of dispersal: interspecific interactions determine spatial dynamics'.

Parasite prevalence is determined by infection state- and risk-dependent dispersal of the host.

The spread of parasites and the emergence of disease are currently threatening global biodiversity and human welfare. To address this threat, we need to better understand those factors that determine parasite persistence and prevalence. It is known that dispersal is central to the spatial dynamics of host-parasite systems. Yet past studies have typically assumed that dispersal is a species-level constant, despite a growing body of empirical evidence that dispersal varies with ecological context, including the risk of infection and aspects of host state such as infection status (parasite-dependent dispersal; PDD). Here, we develop a metapopulation model to understand how different forms of PDD shape the prevalence of a directly transmitted parasite. We show that increasing host dispersal rate can increase, decrease or cause a non-monotonic change in regional parasite prevalence, depending on the type of PDD and characteristics of the host-parasite system (transmission rate, virulence, and dispersal mortality). This result contrasts with previous studies with parasite-independent dispersal which concluded that prevalence increases with host dispersal rate. We argue that accounting for host dispersal responses to parasites is necessary for a complete understanding of host-parasite dynamics and for predicting how parasite prevalence will respond to changes such as human alteration of landscape connectivity. This article is part of the theme issue 'Diversity-dependence of dispersal: interspecific interactions determine spatial dynamics'.

Species interactions affect dispersal: a meta-analysis.

Context-dependent dispersal allows organisms to seek and settle in habitats improving their fitness. Despite the importance of species interactions in determining fitness, a quantitative synthesis of how they affect dispersal is lacking. We present a meta-analysis asking (i) whether the interaction experienced and/or perceived by a focal species (detrimental interaction with predators, competitors, parasites or beneficial interaction with resources, hosts, mutualists) affects its dispersal; and (ii) how the species' ecological and biological background affects the direction and strength of this interaction-dependent dispersal. After a systematic search focusing on actively dispersing species, we extracted 397 effect sizes from 118 empirical studies encompassing 221 species pairs; arthropods were best represented, followed by vertebrates, protists and others. Detrimental species interactions increased the focal species' dispersal (adjusted effect: 0.33 [0.06, 0.60]), while beneficial interactions decreased it (-0.55 [-0.92, -0.17]). The effect depended on the dispersal phase, with detrimental interactors having opposite impacts on emigration and transience. Interaction-dependent dispersal was negatively related to species' interaction strength, and depended on the global community composition, with cues of presence having stronger effects than the presence of the interactor and the ecological complexity of the community. Our work demonstrates the importance of interspecific interactions on dispersal plasticity, with consequences for metacommunity dynamics.This article is part of the theme issue 'Diversity-dependence of dispersal: interspecific interactions determine spatial dynamics'.

Spread of networked populations is determined by the interplay between dispersal behavior and habitat configuration.

Predicting the spread of populations across fragmented habitats is vital if we are to manage their persistence in the long term. We applied network theory with a model and an experiment to show that spread rate is jointly defined by the configuration of habitat networks (i.e., the arrangement and length of connections between habitat fragments) and the movement behavior of individuals. We found that population spread rate in the model was well predicted by algebraic connectivity of the habitat network. A multigeneration experiment with the microarthropod Folsomia candida validated this model prediction. The realized habitat connectivity and spread rate were determined by the interaction between dispersal behavior and habitat configuration, such that the network configurations that facilitated the fastest spread changed depending on the shape of the species' dispersal kernel. Predicting the spread rate of populations in fragmented landscapes requires combining knowledge of species-specific dispersal kernels and the spatial configuration of habitat networks. This information can be used to design landscapes to manage the spread and persistence of species in fragmented habitats.

Negative density-dependent dispersal emerges from the joint evolution of density- and body condition-dependent dispersal strategies.

Empirical studies have documented both positive and negative density-dependent dispersal, yet most theoretical models predict positive density dependence as a mechanism to avoid competition. Several hypotheses have been proposed to explain the occurrence of negative density-dependent dispersal, but few of these have been formally modeled. Here, we developed an individual-based model of the evolution of density-dependent dispersal. This model is novel in that it considers the effects of density on dispersal directly, and indirectly through effects on individual condition. Body condition is determined mechanistically, by having juveniles compete for resources in their natal patch. We found that the evolved dispersal strategy was a steep, increasing function of both density and condition. Interestingly, although populations evolved a positive density-dependent dispersal strategy, the simulated metapopulations exhibited negative density-dependent dispersal. This occurred because of the negative relationship between density and body condition: high density sites produced low-condition individuals that lacked the resources required for dispersal. Our model, therefore, generates the novel hypothesis that observed negative density-dependent dispersal can occur when high density limits the ability of organisms to disperse. We suggest that future studies consider how phenotype is linked to the environment when investigating the evolution of dispersal.

Parasitism Risk and Infection Alter Host Dispersal.

Dispersal determines the spatial dynamics of host-parasite assemblages, particularly during invasions and disease epidemics. The risk of parasitism may create an incentive for dispersal, but infection is expected to reduce dispersal ability, which may alter the host's dispersal response to biotic stressors, including population density. We measured the dispersal of a semiaquatic insect (Notonecta undulata) in aquatic mesocosms in which we manipulated the presence of ectoparasitic Hydrachnidia mites and infected conspecifics. We found that parasitism risk increases host dispersal propensity. Using a flight assay, we determined that parasite infection reduces host dispersal ability. Finally, we used a mark-release-recapture study to investigate the joint effects of both parasitism risk and parasite infection on host dispersal in a natural, spatially structured population. We found that parasitism risk reduced dispersal probability, eliminated positive density-dependent dispersal, and increased dispersal distance. Infection had no effect on dispersal in the natural population. Our results show that parasites can both increase and decrease the movement rates of their hosts, depending on the ecological context, and can alter the host's dispersal response to other biotic stressors. Future studies should consider the consequences of this double-headed impact of parasites for landscape connectivity, population persistence, and host-parasite coevolution.

Phenotype-by-environment interactions influence dispersal.

Numerous studies have demonstrated that dispersal is dependent on both disperser phenotype and the local environment. However, there is substantial variability in the observed strength and direction of phenotype- and environment-dependent dispersal. This has been hypothesized to be the result of interactive effects among the multiple phenotypic and environmental factors that influence dispersal. Here, our goal was to test the hypothesis that these interactions are responsible for generating variation in dispersal behaviour. We achieved these goals by conducting a large, 2-year, mark-release-recapture study of the backswimmer Notonecta undulata in an array of 36 semi-natural ponds. We measured the effects of multiple phenotypic (sex and body size) and environmental (population density and sex ratio) factors, on both dispersal probability and dispersal distance. We found support for the hypothesis that interactive effects influence dispersal and produce variability in phenotype- and environment-dependent dispersal: dispersal probability was dependent on the three-way interaction between sex, body mass and population density. Small males displayed strong, positive density dependence in their dispersal behaviour, while large males and females overall did not respond strongly to density. Small notonectids, regardless of sex, were more likely to disperse, but this effect was strongest at high population densities. Finally, the distance dispersed by backswimmers was a negative function of population density, a pattern which we hypothesize could be related to: (a) individuals from high and low density patches having different dispersal strategies, or (b) the effect of density on dispersal capacity. These results suggest that phenotype-by-environment interactions strongly influence dispersal. Since phenotype- and environment-dependent dispersal has different consequences for ecological and evolutionary dynamics (e.g. metapopulation persistence and local adaptation) than random dispersal, interactive effects may have wide-reaching impacts on populations and communities. We therefore argue that more investment should be made into estimating the effects of multiple, interacting factors on dispersal and determining whether similar interactive effects are acting across systems.

Dispersal depends on body condition and predation risk in the semi-aquatic insect, Notonecta undulata.

Dispersal is the movement of organisms across space, which has important implications for ecological and evolutionary processes, including community composition and gene flow. Previous studies have demonstrated that dispersal is influenced by body condition; however, few studies have been able to separate the effects of body condition from correlated variables such as body size. Moreover, the results of these studies have been inconsistent with respect to the direction of the relationship between condition and dispersal. We examined whether body condition influences dispersal in backswimmers (Notonecta undulata). We also tested whether an interaction between body condition and predation risk (another proximate factor that influences dispersal) could contribute to the previously observed inconsistent relationship between condition and dispersal. We imposed diet treatments on backswimmers in the laboratory, and measured the effects of food availability on body condition and dispersal in the field. We found that dispersal was a positive function of body condition, which may have important consequences for population characteristics such as the rate of gene flow and population growth. However, the effects of body condition and predation risk were additive, not interactive, and therefore, our data do not support the hypothesis that the interaction between condition and predation risk contributes to the inconsistency in the results of previous condition-dependent dispersal studies.

The interactive effects of competition and predation risk on dispersal in an insect.

Dispersal dynamics have significant consequences for ecological and evolutionary processes. Previous work has demonstrated that dispersal can be context-dependent. However, factors affecting dispersal are typically considered in isolation, despite the probability that individuals make dispersal decisions in response to multiple, possibly interacting factors. We examined whether two ecological factors, predation risk and intraspecific competition, have interactive effects on dispersal dynamics. We performed a factorial experiment in mesocosms using backswimmers (Notonecta undulata), flight-capable, semi-aquatic insects. Emigration rates increased with density, and increased with predation risk at intermediate densities; however, predation had minimal effects on emigration at high and low densities. Our results indicate that factorial experiments may be required to understand dispersal dynamics under realistic ecological conditions.