On the evolution of dispersal via heterogeneity in spatial connectivity

Dispersal has long been recognized as a mechanism that shapes many observed ecological and evolutionary processes. Thus, understanding the factors that promote its evolution remains a major goal in evolutionary ecology. Landscape connectivity may mediate the trade-off between the forces in favour of dispersal propensity (e.g. kin-competition, local extinction probability) and those against it (e.g. energetic or survival costs of dispersal). It remains, however, an open question how differing degrees of landscape connectivity may select for different dispersal strategies. We implemented an individual-based model to study the evolution of dispersal on landscapes that differed in the variance of connectivity across patches ranging from networks with all patches equally connected to highly heterogeneous networks. The parthenogenetic individuals dispersed based on a flexible logistic function of local abundance. Our results suggest, all else being equal, that landscapes differing in their connectivity patterns will select for different dispersal strategies and that these strategies confer a long-term fitness advantage to individuals at the regional scale. The strength of the selection will, however, vary across network types, being stronger on heterogeneous landscapes compared with the ones where all patches have equal connectivity. Our findings highlight how landscape connectivity can determine the evolution of dispersal strategies, which in turn affects how we think about important ecological dynamics such as metapopulation persistence and range expansion.     

Evolution of condition-dependent dispersal: A genetic-algorithm search for the ESS reaction norm

Many insects produce two types (winged and wingless) of offspring that greatly differ in dispersal ability. The ratio of the two types often depends on the quality of the local habitat and the crowding experienced by the mother. Here we studied the condition-dependent dispersal that is evolutionarily stable. The model is also applicable to annual plants that produce two types of seeds differing in dispersal rates. The model assumptions are: the population is composed of a number of sites each occupied by a single adult. The total number of offspring produced by a mother depends on the environmental quality of the site that varies over the years and between sites. The ESS fraction of dispersing type as a function of the quality of the habitat (or ESS reaction norm) states that dispersers should not be produced if habitat qualitym is smaller than a critical valuek. Ifm is larger thank, the number of dispersers should increase withm and that of nondispersers should be kept constant. Second, we developed an alternative way of searching for the ESS: the reaction norm is represented as a three-layered neural network, and the parameters (weights and biases) are chosen by genetic algorithm (GA). This method can be extended easily to the cases of multiple environmental factors. There was an optimal (relatively wide) range of mutation rates for weights and biases, outside of which the convergence of the network to the valid ESS was likely to fail. Recombination, or crossing-over, was not effective in improving the success rate. The learned network often shows several characteristic ways of deviation from the ESS. We also examined the case in which the quality of different sites was correlated. In this case the ESS fraction of dispersers increases both with the quality of the site and with the average quality of the whole population in that year.     

The evolution of thermal performance can constrain dispersal during range shifting

Organisms can cope with changing temperature under climate change by either adapting to the temperature at which they perform best and/or by dispersing to more benign locations. The evolution of a new thermal niche during range shifting is, however, expected to be strongly constrained by genetic load because spatial sorting is known to induce fast evolution of dispersal. To broaden our understanding of this interaction, we studied the joint evolution of dispersal and thermal performance curves (TPCs) of a population during range shifting by applying an individual-based spatially explicit model. Always, TPCs adapted to the local thermal conditions. Remarkably, this adaptation coincided with an evolution of dispersal at the shifting range front being equally high or lower than at the trailing edge. This optimal strategy reduces genetic load and highlights that evolutionary dynamics during range shifting change when crucial traits such as dispersal and thermal performance jointly evolve.     

Evolution of positive and negative density-dependent dispersal

Understanding the evolution of density-dependent dispersal strategies has been a major challenge for evolutionary ecologists. Some existing models suggest that selection should favour positive and others negative density-dependence in dispersal. Here, we develop a general model that shows how and why selection may shift from positive to negative density-dependence in response to key ecological factors, in particular the temporal stability of the environment. We find that in temporally stable environments, particularly with low dispersal costs and large group sizes, habitat heterogeneity selects for negative density-dependent dispersal, whereas in temporally variable environments, particularly with high dispersal costs and small group sizes, habitat heterogeneity selects for positive density-dependent dispersal. This shift reflects the changing balance between the greater competition for breeding opportunities in more productive patches, versus the greater long-term value of offspring that establish themselves there, the latter being very sensitive to the temporal stability of the environment. In general, dispersal of individuals out of low-density patches is much more sensitive to habitat heterogeneity than is dispersal out of high-density patches.     

Recruitment trade-offs and the evolution of dispersal mechanisms in plants

In this study we place seed size vs. seed number trade-offs in the context of plant dispersal ability. The objective was to suggest explanations for the evolution of different seed dispersal mechanisms, in particular fleshy fruits, wind dispersal and the maintenance of unassisted dispersal. We suggest that selection for improved dispersal may act either by increasing the intercept of a dispersal curve (log seed number vs. distance) or by flattening the slope of the curve. 'Improved dispersal' is defined as a marginal increase in the number of recruits sited at some (arbitrary) distance away from the parent plant. Increasing the intercept of the dispersal curve, i.e. producing more seeds, is associated with a reduction in seed size, which in turn affects the recruitment ability, provided that this ability is related to seed size. If recruitment is related to seed size there will be a recruitment cost of evolving increased seed production. On the other hand, a flattening of the slope by evolving dispersal attributes is likely to be associated with a fecundity cost. An exception is wind dispersal where smaller (and hence more numerous) seeds may lead to more efficient dispersal. We derive two main predictions: If recruitment is strongly related to seed size, selection for improved dispersal acts on the slope of the dispersal curve, i.e. by favouring evolution of dispersal attributes on seeds or fruits. If, on the other hand, recruitment is only weakly related to seed size (or not related, or negatively related), selection for improved dispersal favours increased seed production. Despite its simplicity, the model suggests explanations for (i) why so many plant species lack special seed dispersal attributes, (ii) differences in dispersal spectra among plant communities, and (iii) adaptive radiation in seed size and dispersal attributes during angiosperm evolution. This revised version was published online in July 2006 with corrections to the Cover Date.     

On several conjectures from evolution of dispersal

We address several conjectures raised in Cantrell et al. [Evolution of dispersal and ideal free distribution, Math. Biosci. Eng. 7 (2010), pp. 17–36 [9 Cantrell, R. S., Cosner, C. and Lou, Y. 2010. Evolution of dispersal and ideal free distribution. Math. Biosci. Eng., 7: 17–36. [Crossref], [PubMed], [Web of Science ®] , [Google Scholar]]] concerning the dynamics of a diffusion–advection–competition model for two competing species. A conditional dispersal strategy, which results in the ideal free distribution of a single population at equilibrium, was found in Cantrell et al. [9 Cantrell, R. S., Cosner, C. and Lou, Y. 2010. Evolution of dispersal and ideal free distribution. Math. Biosci. Eng., 7: 17–36. [Crossref], [PubMed], [Web of Science ®] , [Google Scholar]]. It was shown in [9 Cantrell, R. S., Cosner, C. and Lou, Y. 2010. Evolution of dispersal and ideal free distribution. Math. Biosci. Eng., 7: 17–36. [Crossref], [PubMed], [Web of Science ®] , [Google Scholar]] that this special dispersal strategy is a local evolutionarily stable strategy (ESS) when the random diffusion rates of the two species are equal, and here we show that it is a global ESS for arbitrary random diffusion rates. The conditions in [9 Cantrell, R. S., Cosner, C. and Lou, Y. 2010. Evolution of dispersal and ideal free distribution. Math. Biosci. Eng., 7: 17–36. [Crossref], [PubMed], [Web of Science ®] , [Google Scholar]] for the coexistence of two species are substantially improved. Finally, we show that this special dispersal strategy is not globally convergent stable for certain resource functions, in contrast with the result from [9 Cantrell, R. S., Cosner, C. and Lou, Y. 2010. Evolution of dispersal and ideal free distribution. Math. Biosci. Eng., 7: 17–36. [Crossref], [PubMed], [Web of Science ®] , [Google Scholar]], which roughly says that this dispersal strategy is globally convergent stable for any monotone resource function.     

Accelerating invasion rates result from the evolution of density-dependent dispersal

Evolutionary processes play an important role in shaping the dynamics of range expansions, and selection on dispersal propensity has been demonstrated to accelerate rates of advance. Previous theory has considered only the evolution of unconditional dispersal rates, but dispersal is often more complex. For example, many species emigrate in response to crowding. Here, we use an individual-based model to investigate the evolution of density dependent dispersal into empty habitat, such as during an invasion. The landscape is represented as a lattice and dispersal between populations follows a stepping-stone pattern. Individuals carry three ‘genes’ that determine their dispersal strategy when experiencing different population densities. For a stationary range we obtain results consistent with previous theoretical studies: few individuals emigrate from patches that are below equilibrium density. However, during the range expansion of a previously stationary population, we observe evolution towards dispersal strategies where considerable emigration occurs well below equilibrium density. This is true even for moderate costs to dispersal, and always results in accelerating rates of range expansion. Importantly, the evolution we observe at an expanding front depends upon fitness integrated over several generations and cannot be predicted by a consideration of lifetime reproductive success alone. We argue that a better understanding of the role of density dependent dispersal, and its evolution, in driving population dynamics is required especially within the context of range expansions.     

Phenotypic evolution in stochastic environments: The contribution of frequency- and density-dependent selection

Understanding how environmental variation affects phenotypic evolution requires models based on ecologically realistic assumptions that include variation in population size and specific mechanisms by which environmental fluctuations affect selection. Here we generalize quantitative genetic theory for environmentally induced stochastic selection to include general forms of frequency- and density-dependent selection. We show how the relevant fitness measure under stochastic selection relates to Fisher's fundamental theorem of natural selection, and present a general class of models in which density regulation acts through total use of resources rather than just population size. In this model, there is a constant adaptive topography for expected evolution, and the function maximized in the long run is the expected factor restricting population growth. This allows us to generalize several previous results and to explain why apparently “-selected” species with slow life histories often have low carrying capacities. Our joint analysis of density- and frequency-dependent selection reveals more clearly the relationship between population dynamics and phenotypic evolution, enabling a broader range of eco-evolutionary analyses of some of the most interesting problems in evolution in the face of environmental variation.     

Negative density-dependent dispersal in the American black bear (Ursus americanus) revealed by noninvasive sampling and genotyping

Although the dispersal of animals is influenced by a variety of factors, few studies have used a condition-dependent approach to assess it. The mechanisms underlying dispersal are thus poorly known in many species, especially in large mammals. We used 10 microsatellite loci to examine population density effects on sex-specific dispersal behavior in the American black bear, Ursus americanus. We tested whether dispersal increases with population density in both sexes. Fine-scale genetic structure was investigated in each of four sampling areas using Mantel tests and spatial autocorrelation analyses. Our results revealed male-biased dispersal pattern in low-density areas. As population density increased, females appeared to exhibit philopatry at smaller scales. Fine-scale genetic structure for males at higher densities may indicate reduced dispersal distances and delayed dispersal by subadults.     

The color of noise and the evolution of dispersal

The process of dispersal is vital for the long-term persistence of all species and hence is a ubiquitous characteristic of living organisms. A present challenge is to increase our understanding of the factors that govern the dispersal rate of individuals. Here I extend previous work by incorporating both spatial and temporal heterogeneity in terms of patch quality into a spatially explicit lattice model. The spatial heterogeneity is modeled as a two-dimensional fractal landscape, while temporal heterogeneity is included by using one-dimensional noise. It was found that the color of both the spatial and temporal variability influences the rate of dispersal selected as reddening of the temporal noise leads to a reduction in dispersal, while reddening of spatial variability results in an increase in the dispersal rate. These results demonstrate that the color of environmental noise should be considered in future studies looking at the evolution of life history characteristics.     

How landscapes affect the evolution of dispersal behaviour in reef fishes: results from an individual‐based model

The vast majority of tropical reef fishes have a sedentary adult phase and pelagic larval phase that is potentially highly dispersive. Dispersal may be favoured by a wide range of factors including the arrangement of suitable habitat in space. In this paper the dispersal strategy of individuals is followed and allowed to evolve in a simplified model of three different landscapes: an enclosed sea, an open archipelago and a barrier reef. The three landscapes have very different characteristics, but all have similar spatial clumping of reef habitat. In all landscapes, as minimum time to settlement increases, evolved movement strategy also increases and longer settlement windows favour dispersal. In the archipelago movement is not maximized until the minimum pelagic duration is longer than in the other landscapes. The model predicts that, given the same pelagic duration, species from enclosed seas should have more dispersive behaviours than those from open archipelagos, because of the density of habitat and the aggregation of habitat in space affect the likelihood of larvae finding suitable habitat for settlement.     

Conditional dispersal, clines, and the evolution of dispersiveness

Conditional dispersal, in which an individual’s decision over whether to disperse is a response to environmental conditions, features prominently in studies of dispersal evolution. Using models of clines, I examine how one widely discussed cost of dispersal, namely, that dispersal impedes local adaptation, changes with conditional dispersal and what this implies for dispersal evolution. I examine the consequences for dispersal evolution of the responsiveness of dispersal to the environment, the accuracy of any proximal cues that individuals rely upon to assess habitat quality, and whether dispersal responds to fitness itself or only to some fitness components (juvenile survivorship). All of the conditional dispersal behaviors that I consider weaken the indirect cost of dispersal inhibiting local adaptation. However, if individuals rely on imprecise cues to assess habitat quality and base dispersal decisions on juvenile survivorship, then conditional dispersal can incur additional costs by exacerbating overcrowding. Conditional dispersal initially leads to steeper clines in traits under direct selection, but when dispersiveness can itself evolve, conditional dispersal allows sigmoidal clines to persist long after those obtained with unconditional movement would become stepped. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Monitoring and modeling density-dependent growth of anchorage-dependent cells

The density-dependent growth of Chinese hamster ovary (CHO) cells was monitored on-line by using an inverted microscope. A flow system was employed for cell cultivation so that nutrient concentration could be maintained and metabolic wastes were removed. With the help of video image analysis, local cells density could be accurately calculated and cell motility and exposed cell surface area could be estimated. A computer program which accounted for change of sell size and translocation of cells was developed to stimulate cell growth. The stimulated results of the population dynamics and the variations in cell size showed good agreement with our experimental observations, Cell motility and initial cell distribution on the substratum were found to have strong effect on cell growth.     

Joint evolution of sex ratio and dispersal: conditions for higher dispersal rates from good habitats

We analyze models of evolution of sex ratio conditional on habitat quality and with sex specific dispersal. Previous analysis concluded that the main constraint on sex ratio is habitat choice and leads to overproduction of the most dispersing sex in low quality habitat. Here, we analyze three models with finite local populations and show that constraints on sex ratio can balance constraints on habitat choice. In the first model, dispersal rates are fixed. In the second, sex specific dispersal can evolve independently of the habitat quality. These models suggests that sex ratio evolution can lead to higher global dispersal rates (mean of male and female dispersal rates) from high quality habitats. In the last model dispersal evolves conditionally with both sex and habitat. Our models suggests that conditions for overproduction of the most dispersing sex in high quality habitat are frequent. The predictions of the models with evolving dispersal contrast with patterns generally described in nature. We discuss possible reasons of this difference.     

Joint evolution of dispersal and connectivity

Functional connectivity, the realized flow of individuals between the suitable sites of a heterogeneous landscape, is a prime determinant of the maintenance and evolution of populations in fragmented habitats. While a large body of literature examines the evolution of dispersal propensity, it is less known how evolution shapes functional connectivity via traits that influence the distribution of the dispersers. Here, we use a simple model to demonstrate that, in a heterogeneous environment with clustered and solitary sites (i.e., with variable structural connectivity), the evolutionarily stable population contains strains that are strongly differentiated in their pattern of connectivity (local vs. global dispersal), but not necessarily in the fraction of dispersed individuals. Also during evolutionary branching, selection is disruptive predominantly on the pattern of connectivity rather than on dispersal propensity itself. Our model predicts diversification along a hitherto neglected axis of dispersal strategies and highlights the role of the solitary sites—the more isolated and therefore seemingly less important patches of habitat—in maintaining global dispersal that keeps all sites connected.     

Distribution patterns and dispersal strategies of Galiciales

In Caliciales both passive and active spore dispersal occurs. A review of the distribution of 162 species is given. Species with small spores have wider distribution than species with large spores, irrespective of dispersal strategy. In antitropical species there is only rarely a differentiation between Northern and Southern Hemisphere populations. It is concluded that in several small-spored genera long-distance dispersal has been instrumental in dispersing the species and range extensions have been much more frequent than in large-spored species for which vicariance explanations are more appropriate. Comparisons with bryophyte distributions show differences in dispersal strategies and distribution patterns.