Density-dependent Dispersal Research Articles

Landscape properties and density dependence shape the movement patterns of three threatened butterflies

ContextConservation of endangered species necessitates an in-depth understanding of their ecological requirements. Particularly in landscape ecology, the behavioural tendencies of threatened butterfly species in Gotland, a biodiversity-rich island in the Baltic Sea, become crucial.ObjectivesThe primary aim of this study was to elucidate the movement patterns of three threatened butterfly species—Euphydryas aurinia, Parnassius apollo, and Phengaris arion—in Gotland and to identify the influence of specific land characteristics on these patterns.MethodsOur study, conducted from 2017 to 2020 across 60 km2 in Gotland, involved detailed capture-mark-recapture (CMR) efforts of 29,584 captures including 16,223 unique butterflies. We investigate the departure and arrival events of butterflies, specifically focusing on the associations between movements when individuals leave or enter a hectare grid different from their previously recorded location and key landscape features: open vegetated land, ground moisture, and forest cover. We model landscape features to examine the interplay between these and butterfly movement patterns, providing insights into preferred landscape features and conservation strategies.ResultsAmong the 4821 arrivals and 5083 departures documented, the species exhibited differential responses to the evaluated habitat features. Both E. aurinia and P. apollo displayed a positive density-dependent dispersal, while P. arion’s movements were not significantly associated with any of the examined habitat features. Landscape properties like open vegetated land and ground moisture index statistically influenced the likelihood of arrival and departure.ConclusionsThe study accentuates the relationship between land cover and the behavioural tendencies of the subject butterfly species. It has broader implications for the targeted habitat management strategies that would benefit threatened butterfly populations in Gotland.

Age and Density of Mated Females Affect Dispersal Strategies in Spider Mite Tetranychus ludeni Zacher.

The dispersal strategies of a species can affect its invasion success. Investigations into the dispersal strategies of invasive species in relation to different factors help improve our understanding of invasion mechanisms and provide knowledge for population management and invasion evaluation. Tetranychus ludeni Zacher (Acari: Tetranychidae) is an invasive species which is native to Europe but is now cosmopolitan. Here, we examined the effects of age and density on dispersal in mated females. Our results show that older females that are capable of producing more eggs within 24 h were more likely to disperse and moved longer distances than younger ones with fewer eggs. Older females spread most of their eggs out of their natal habitats and over longer distances, which reduced competition and increased offspring fitness. Females exhibited significantly increased dispersal probability and distances with an increase in population density to avoid crowding. The synchronization of dispersal and reproduction, along with the positive density-dependent dispersal strategy, may facilitate the habitat colonization and invasion speed of T. ludeni.

Tipping points emerge from weak mutualism in metacommunities.

The coexistence of obligate mutualists is often precariously close to tipping points where small environmental changes can drive catastrophic shifts in species composition. For example, microbial ecosystems can collapse by the decline of a strain that provides an essential resource on which other strains cross-feed. Here, we show that tipping points, ecosystem collapse, bistability and hysteresis arise even with very weak (non-obligate) mutualism provided the population is spatially structured. Based on numeric solutions of a metacommunity model and mean-field analyses, we demonstrate that weak mutualism lowers the minimal dispersal rate necessary to avoid stochastic extinction, while species need to overcome a mean threshold density to survive in this low dispersal rate regime. Our results allow us to make numerous predictions for mutualistic metacommunities regarding tipping points, hysteresis effects, and recovery from external perturbations, and let us draw general conclusions for ecosystems even with random, not necessarily mutualistic, interactions and systems with density-dependent dispersal rather than direct mutualistic interactions.

Large-scale genotypic identification reveals density-dependent natal dispersal patterns in an elusive bird of prey

BackgroundNatal dispersal, the distance between site of birth and site of first breeding, has a fundamental role in population dynamics and species’ responses to environmental changes. Population density is considered a key driver of natal dispersal. However, few studies have been able to examine densities at both the natal and the settlement site, which is critical for understanding the role of density in dispersal. Additionally, the role of density on natal dispersal remains poorly understood in long-lived and slowly reproducing species, due to their prolonged dispersal periods and often elusive nature. We studied the natal dispersal of the white-tailed eagle (Haliaeetus albicilla) in response to local breeder densities. We investigated the effects of the number of active territories around the natal site on (a) natal dispersal distance and (b) the difference between natal and settlement site breeder density. We were interested in whether eagles showed tendencies of conspecific attraction (positive density-dependence) or intraspecific competition (negative density-dependence) and how this related to settlement site breeder density.MethodsWe used a combination of long-term visual and genotypic identification to match individuals from their breeding site to their natal nest. We identified natal dispersal events for 355 individuals hatched between 1984 and 2015 in the Baltic Sea coast and Arctic areas of Finland. Of those, 251 were identified by their genotype.ResultsIndividuals born in high-density areas dispersed shorter distances than those born in low-density areas, but settled at lower density breeding sites in comparison to their natal site. Eagles born in low natal area densities dispersed farther but settled in higher density breeding sites compared to their natal site.ConclusionsWe show that eagles might be attracted by conspecifics (positive density-dependence) to identify high-quality habitats or find mates, but do not settle in the most densely populated areas. This indicates that natal dispersal is affected by an interplay of conspecific attraction and intraspecific competition, which has implications for population dynamics of white-tailed eagles, but also other top predators. Furthermore, our study demonstrates the value of long-term collection of both nestling and (non-invasive) adult DNA samples, and thereafter using genotype matching to identify individuals in long-lived and elusive species.

Regime shifts, extinctions and long transients in models of population dynamics with density-dependent dispersal

Predicting extinctions resulting from ecosystems' regime shifts has long been a focus of biological conservation and ecological management. Mathematical modelling plays a key role in assessing the possibility of such events. Traditionally, however, models focused on long-term, asymptotic behaviour of ecosystems. Meanwhile, the environment is usually non-stationary, which may mean that the long-term behaviour is never observed. Correspondingly, over the last two decades there has been a growing appreciation of the role of transients both in empirical ecology and theoretical studies, in particular in the context of species extinctions. In this paper, we theoretically explore long transients and extinctions occurring in several paradigmatic models of increasing complexity, such as single species, two-species and three-species systems. We consider the population dynamics in a local ‘patch’, the patch being connected to the rest of the population via density-dependent dispersal. We consider both deterministic and stochastic scenarios. We discover many different patterns of long transient dynamics with quick regime shifts between ‘safe’ (persistent) dynamics and unsafe ones resulting in extinctions. Remarkably, delayed extinction can occur after thousands of generations of apparently safe population dynamics. We classify transient regimes and reveal their underlying mechanisms. Environmental noise can either shorten transients or can create a new type of a long transient. Our study suggests that not only are long transients ubiquitous, but there is also a great variety of them. The omnipresence of long transients emphasizes the need to account for them in nature conservation programs as well as future theoretical research.

Total equilibrium biomass of a two-patch logistic equation with density-dependent dispersal

Total equilibrium biomass of a two-patch logistic equation with density-dependent dispersal

Landscape connectivity alters the evolution of density-dependent dispersal during pushed range expansions

Landscape connectivity alters the evolution of density-dependent dispersal during pushed range expansions

Providing evidence for the conservation of a rare forest butterfly: Results from a three-year capture-mark-recapture study

Species living in spatially structured populations require a network of interconnected habitat patches. Due to changes in forest management, this network of habitat patches has been lost for insect species inhabiting open spots within forests. We studied two of the last populations of the Southern White Admiral (Limenitis reducta) in Germany. The aim of our study was to provide information for the conservation of this species. We conducted a capture-mark-recapture study over three consecutive years and we estimated population sizes and demographic parameters using Jolly-Seber and Cormack-Jolly-Seber models. Furthermore, we used different dispersal kernels to study the dispersal of L. reducta. We found that apparent survival rates differed greatly between the sexes. The mean residence times were eight to nine days for males and only two to four days for females. Apparent survival rates of both sexes decreased with increasing wing deterioration. Total population sizes of L. reducta varied between sites and years and ranged between 61 and 123. Daily abundances were generally low, especially that of females. The mean dispersal distance of individual recaptured butterflies increased in years with higher population densities, suggesting density-dependent dispersal. The dispersal data in our study was clearly bimodal, probably reflecting ‘routine movement’ at short distances and ‘displacement movement’ between habitat patches at longer distances. Consequently, the processes generating the dispersal data were better represented by the lognormal mixture model than by the negative exponential and the inverse power function. The mixed kernel predicted that about 9 % of the population disperses over > 1 km but that long-distance dispersal is rare. Our study highlights the urgent need for conservation measures to protect L. reducta in Germany and, based on our data, we recommend to create new habitat patches at distances of 1 to 1.5 km from existing habitats.

Cover Picture and Issue Information

Functional EcologyVolume 37, Issue 3 p. 473-475 COVER PICTURE AND ISSUE INFORMATIONFree Access Cover Picture and Issue Information First published: 02 March 2023 https://doi.org/10.1111/1365-2435.14073AboutPDF ToolsRequest permissionExport citationAdd to favoritesTrack citation ShareShare Give accessShare full text accessShare full-text accessPlease review our Terms and Conditions of Use and check box below to share full-text version of article.I have read and accept the Wiley Online Library Terms and Conditions of UseShareable LinkUse the link below to share a full-text version of this article with your friends and colleagues. Learn more.Copy URL Share a linkShare onFacebookTwitterLinkedInRedditWechat Graphical Abstract Flowering plants need pollinators to transfer pollen, such as those pictured on this Tamarix chinensis (Credit: Le Jiao). The movement behavior and strategy of pollinators significantly affects pollination success and the subsequent seed set, especially for plant populations that are faced with habitat loss. Jiao et al. 2022 (DOI: 10.1111/1365-2435.14252) simulated two key stages of pollinator movement during the pollination process: a foraging response stage towards plant patches mediated by floral scents, and an individual dispersal stage between plant patches mediated by a densitydependent dispersal strategy. Plant reproduction was negatively affected by habitat loss, as the emission amount of floral scents and the foraging success rate of pollinators decreased in the context of habitat loss. However, the pollinator movement strategy (i.e. density-dependent dispersal strategy) can help mitigate adverse impacts by promoting visitation to suboptimal plant patches, thereby improving the persistence of the plant population in fragmented landscapes. Altogether, this simulation result points to the positive role of pollinator movement strategy in alleviating the negative effects of habitat loss on plant reproduction. Volume37, Issue3March 2023Pages 473-475 RelatedInformation

Density‐dependent dispersal strategy of pollinator moderates the adverse effect of habitat loss on plant reproduction: An integrated model based on pollinators' behavioural response

Abstract Major challenges for plant conservation are predicting the effect of habitat loss on pollination success and plant reproduction potential. Most studies report that pollinator movement is affected by quantitative and spatial characteristics of landscapes. However, little is known about the role of pollinator movement, impacted by floral volatiles and intraspecies interaction, on plant reproduction in fragmented landscapes. To clarify the effect of pollinator movement on plant reproduction relative to habitat loss, we developed an integrated model incorporating pollinator's foraging response with its dispersal process mediated by a density‐dependent dispersal (DDD) strategy. This model performed better in capturing behaviour response of pollinators than do current methods. The integrated model was verified with field results of pollinator visitation and plant reproduction of saltcedar Tamarix chinensis inhabiting the Yellow River Delta, and then was compared against a dispersal strategy called density‐independent dispersal (DID). The model was applied to landscapes with various non‐habitat percentage (NHP) to explore the effect of habitat loss on plant reproduction. Results suggested that saltcedar populations differ in their responses to habitat loss, which depended on the spatial scales considered. At landscape scale, increasing NHP significantly inhibited dispersion extent of floral volatiles and therefore reduced pollinator visitation and subsequent seed production, especially when NHP exceeded the critical threshold of 0.6. However, at patch scale, comparing with DID strategy, the DDD strategy enabled pollinators to increase their utilization of flowers by 43.42% and 6.79% in low‐density and distant plant patches, whereas their utilization was reduced by 7.75% and 2.24% in high‐density and central patches, respectively. Plant reproduction was improved correspondingly in low‐density and distant patches under different NHPs. Consequently, habitat loss inhibits the volatile dispersion and interferes the foraging success of pollinators, a major factor influencing plant reproduction at landscape scale. At patch scale, adaptive utilization of pollinators exhibiting DDD strategy alleviates the negative effect of habitat loss on plant production, and maintain plant population persistence. Since pollinator behavioural response is critical to plant reproduction, we recommend the use of the here presented integrated model to assess the impact of habitat loss on plant reproduction. Read the free Plain Language Summary for this article on the Journal blog.

A simulation test of the prediction that density-dependent dispersal promotes female-biased sex allocation in viscous populations.

A classic result of sex-allocation theory is that the sex ratio is predicted to be invariant with respect to the rate of dispersal. However, a recent mathematical analysis has suggested that if individuals are able to adjust their probability of dispersal according to the local density of their neighbourhood, then a lower rate of dispersal will be associated with greater female-bias. Here, we perform a computer simulation test of this prediction. Our simulation data provide strong qualitative support for the prediction, and a Monte Carlo randomization test of significance allows us to reject the null hypothesis of the invariance relationship.

To go or not to go: variable density-dependent dispersal in small mammals

AbstractPopulation density has been widely understood to be a key influencer of dispersal behavior; however, the generality of density-dependent (DD) dispersal in vertebrates is unclear. We conducted a review of the available empirical data on small mammal DD dispersal, distinguishing between the three dispersal stages: emigration, immigration, and transience (dispersal distance). We focused on small mammals because they are a well-studied, functionally similar group of vertebrates, with a distinct ecological importance. We also examined the effect of season, body mass, study length, and study type on the strength and direction of DD dispersal. The majority of emigration and dispersal distance studies reported negative density dependence, while immigration was mostly independent of density. No correlative patterns were detected; however, interpretation of the available data was hindered by inconsistencies in experimental and analytical approach across studies. Our results suggest that the three phases of the dispersal process may be influenced differently by density and highlight the importance of distinguishing between emigration, immigration, and transience when considering the effects of density dependence. As well, our study identifies several limitations with the current available data which limit the ability to compare DD dispersal behavior across systems, and calls for future investigations that consider all three phases of dispersal in the same system.

Plant-host shift, spatial persistence, and the viability of an invasive insect population

Assessing the effects of a plant-host shift is important for monitoring insect populations over long time periods and for interventions in a conservation or pest management framework. In a heterogeneous environment, individuals may disperse between sources and sinks in order to persist. Here we propose a single-species two-patch model that aims to capture the generational movement of an insect that exhibits density-dependent dispersal, to see how shifting between hosts could alter its viability and asymptotic dynamics. We then analyse the stability and persistence properties of the model and further validate it using parameter estimates derived from laboratory experiments. In order to evaluate the potential of this model, we applied it to Drosophila suzukii (Diptera: Drosophilidae), which has become a harmful pest in several countries around the world. Although many studies have investigated the preference and attractiveness of potential hosts on this invasive drosophilid, no studies thus far have investigated whether a shift of fruit host could affect such a species’ ecological viability or spatiotemporal persistence. The model results show that a shift in host choice can significantly affect the growth potential and fecundity of a species such as D. suzukii, which ultimately could aid such invasive populations in their ability to persist within a changing environment.

Density-dependent dispersal promotes female-biased sex allocation in viscous populations

A surprising result emerging from the theory of sex allocation is that the optimal sex ratio is predicted to be completely independent of the rate of dispersal. This striking invariance result has stimulated a huge amount of theoretical and empirical attention in the social evolution literature. However, this sex-allocation invariant has been derived under the assumption that an individual's dispersal behaviour is not modulated by population density. Here, we investigate how density-dependent dispersal shapes patterns of sex allocation in a viscous-population setting. Specifically, we find that if individuals are able to adjust their dispersal behaviour according to local population density, then they are favoured to do so, and this drives the evolution of female-biased sex allocation. This result obtains because, whereas under density-independent dispersal, population viscosity is associated not only with higher relatedness—which promotes female bias—but also with higher kin competition—which inhibits female bias—under density-dependent dispersal, the kin-competition consequences of a female-biased sex ratio are entirely abolished. We derive analytical results for the full range of group sizes and costs of dispersal, under haploid, diploid and haplodiploid modes of inheritance. These results show that population viscosity promotes female-biased sex ratios in the context of density-dependent dispersal.

Synchrony and Stability in Trophic Metacommunities: When Top Predators Navigate in a Heterogeneous World

Ecosystem stability strongly depends on spatial aspects since localized perturbations spread across an entire region through species dispersal. Assessing the synchrony of the response of connected populations is fundamental to understand stability at different scales because if populations fluctuate asynchronously, the risk of their simultaneous extinction is low, thus reducing the species' regional extinction risk. Here, we consider a metacommunity model consisting of two food chains connected by dispersal and we review the various mechanisms governing the transmission of small perturbations affecting populations in the vicinity of equilibrium. First, we describe how perturbations propagate vertically (i.e., within food chains through trophic interactions) and horizontally (i.e., between food chains through dispersal) in metacommunities. Then, we discuss the mechanisms susceptible to alter synchrony patterns such as density-depend dispersal or spatial heterogeneity. Density-dependent dispersal, which is the influence of prey or predator abundance on dispersal, has a major impact because the species with the highest coefficient of variation of biomass governs the dispersal rate of the dispersing species and determines the synchrony of its populations, thus bypassing the classic vertical transmission of perturbations. Spatial heterogeneity, which is a disparity between patches of the attack rate of predators on prey in our model, alters the vertical transmission of perturbations in each patch, thus making synchrony dependent on which patch is perturbed. Finally, by combining our understanding of the impact of each of these mechanisms on synchrony, we are able to full explain the response of realistic metacommunities such as the model developed by Rooney et al. (2006). By disentangling the main mechanisms governing synchrony, our metacommunity model provides a broad insight into the consequences of spacial aspects on food web stability.

Population viscosity promotes altruism under density-dependent dispersal.

A basic mechanism of kin selection is population viscosity, whereby individuals do not move far from their place of birth and hence tend to be surrounded by relatives. In such circumstances, even indiscriminate altruism among neighbours will often involve interactions between kin, which has a promoting effect on the evolution of altruism. This has the potential to explain altruistic behaviour across the whole tree of life, including in taxa for which recognition of kin is implausible. However, population viscosity may also intensify resource competition among kin, which has an inhibitory effect on altruism. Indeed, in the simplest scenario, in which individuals disperse with a fixed probability, these two effects have been shown to exactly cancel such that there is no net impact of viscosity on altruism. Here, we show that if individuals are able to disperse conditionally upon local density, they are favoured to do so, with more altruistic neighbourhoods exhibiting a higher rate of dispersal and concomitant relaxation of kin competition. Comparing across different populations or species, this leads to a negative correlation between overall levels of dispersal and altruism. We demonstrate both analytically and using individual-based simulations that population viscosity promotes the evolution of altruism under density-dependent dispersal.

When higher carrying capacities lead to faster propagation

Finding general patterns in the expansion of natural populations is a major challenge in ecology and invasion biology. Classical spatio-temporal models predict that the carrying capacity (K) of the environment should have no influence on the speed (v) of an expanding population. We tested the generality of this statement with reaction-diffusion equations, stochastic individual-based models, and microcosms experiments with Trichogramma chilonis wasps. We investigated the dependence between K and v under different assumptions: null model (Fisher-KPP-like assumptions), strong Allee effects, and positive density-dependent dispersal. These approaches led to similar and complementary results. Strong Allee effects, positive density-dependent dispersal and demographic stochasticity in small populations lead to a positive dependence between K and v. A positive correlation between carrying capacity and propagation speed might be more frequent than previously expected, and be the rule when individuals at the edge of a population range are not able to fully drive the expansion.

Examining the effects of heterospecific abundance on dispersal in forest small mammals

AbstractThe effects of conspecific densities on dispersal have been well documented. However, while positive and negative density-dependent dispersal based on conspecific densities often are shown to be the result of intraspecific competition or facilitation, respectively, the effects of heterospecific densities on dispersal have been examined far less frequently. This gap in knowledge warrants investigation given the potential for the analogous processes of interspecific competition and heterospecific attraction to influence dispersal patterns and behavior. Here we use a long-term live-trapping study of deer mouse (Peromyscus maniculatus), eastern chipmunk (Tamias striatus), red-backed vole (Myodes gapperi), and jumping mice (Napaeozapus insignis and Zapus hudsonius) to examine the effects of variation in conspecific and heterospecific abundances on dispersal frequency. In terms of conspecific abundance, jumping mice were more likely to disperse from areas with fewer conspecifics, while red-backed voles and chipmunks did not respond to variation in conspecific abundances in their dispersal frequencies. While there were no statistically significant effects of variation in heterospecific abundances on dispersal frequency, some effect sizes for heterospecific abundance effects on dispersal met or exceeded those of conspecific abundances. Conspecific abundances clearly can affect dispersal by some species in this system, but the effects of heterospecific abundances on dispersal frequency are less clear. Based on effect sizes, it appears that there may be potential for heterospecific effects on dispersal by some species in the community, although the strength and causes of these relationships remain unclear.

Virus-Host Interactions Shape Viral Dispersal Giving Rise to Distinct Classes of Traveling Waves in Spatial Expansions

Reaction-diffusion waves have long been used to describe the growth and spread of populations undergoing a spatial range expansion. Such waves are generally classed as either pulled, where the dynamics are driven by the very tip of the front and stochastic fluctuations are high, or pushed, where cooperation in growth or dispersal results in a bulk-driven wave in which fluctuations are suppressed. These concepts have been well studied experimentally in populations where the cooperation leads to a density-dependent growth rate. By contrast, relatively little is known about experimental populations that exhibit density-dependent dispersal. Using bacteriophage T7 as a test organism, we present novel experimental measurements that demonstrate that the diffusion of phage T7, in a lawn of host E. coli, is hindered by steric interactions with host bacteria cells. The coupling between host density, phage dispersal and cell lysis caused by viral infection results in an effective density-dependent diffusion coefficient akin to cooperative behavior. Using a system of reaction-diffusion equations, we show that this effect can result in a transition from a pulled to pushed expansion. Moreover, we find that a second, independent density-dependent effect on phage dispersal spontaneously emerges as a result of the viral incubation period, during which phage is trapped inside the host unable to disperse. Additional stochastic agent-based simulations reveal that lysis time dramatically affects the rate of diversity loss in viral expansions. Taken together, our results indicate both that bacteriophage can be used as a controllable laboratory population to investigate the impact of density-dependent dispersal on evolution, and that the genetic diversity and adaptability of expanding viral populations could be much greater than is currently assumed.

Negative density-dependent dispersal in tsetse (Glossina spp): An artefact of inappropriate analysis.

Published analysis of genetic material from field-collected tsetse (Glossina spp, primarily from the Palpalis group) has been used to predict that the distance (δ) dispersed per generation increases as effective population densities (De) decrease, displaying negative density-dependent dispersal (NDDD). Using the published data we show this result is an artefact arising primarily from errors in estimates of S, the area occupied by a subpopulation, and thereby in De. The errors arise from the assumption that S can be estimated as the area () regarded as being covered by traps. We use modelling to show that such errors result in anomalously high correlations between and and the appearance of NDDD, with a slope of -0.5 for the regressions of log() on log(), even in simulations where we specifically assume density-independent dispersal (DID). A complementary mathematical analysis confirms our findings. Modelling of field results shows, similarly, that the false signal of NDDD can be produced by varying trap deployment patterns. Errors in the estimates of δ in the published analysis were magnified because variation in estimates of S were greater than for all other variables measured, and accounted for the greatest proportion of variation in . Errors in census population estimates result from an erroneous understanding of the relationship between trap placement and expected tsetse catch, exacerbated through failure to adjust for variations in trapping intensity, trap performance, and in capture probabilities between geographical situations and between tsetse species. Claims of support in the literature for NDDD are spurious. There is no suggested explanation for how NDDD might have evolved. We reject the NDDD hypothesis and caution that the idea should not be allowed to influence policy on tsetse and trypanosomiasis control.