Dispersal strategies, few dominating or many coexisting: the effect of environmental spatial structure and multiple sources of mortality

Interspecific competition, life history traits, environmental heterogeneity and spatial structure as well as disturbance are known to impact the successful dispersal strategies in metacommunities. However, studies on the direction of impact of those factors on dispersal have yielded contradictory results and often considered only few competing dispersal strategies at the same time. We used a unifying modeling approach to contrast the combined effects of species traits (adult survival, specialization), environmental heterogeneity and structure (spatial autocorrelation, habitat availability) and disturbance on the selected, maintained and coexisting dispersal strategies in heterogeneous metacommunities. Using a negative exponential dispersal kernel, we allowed for variation of both species dispersal distance and dispersal rate. We showed that strong disturbance promotes species with high dispersal abilities, while low local adult survival and habitat availability select against them. Spatial autocorrelation favors species with higher dispersal ability when adult survival and disturbance rate are low, and selects against them in the opposite situation. Interestingly, several dispersal strategies coexist when disturbance and adult survival act in opposition, as for example when strong disturbance regime favors species with high dispersal abilities while low adult survival selects species with low dispersal. Our results unify apparently contradictory previous results and demonstrate that spatial structure, disturbance and adult survival determine the success and diversity of coexisting dispersal strategies in competing metacommunities.     

Diversity and Disturbances in the Antarctic Megabenthos: Feasible versus Theoretical Disturbance Ranges

The intermediate disturbance hypothesis (IDH) predicts a hump-shaped relationship between regional diversity and the disturbance rate. We tested the IDH for the megabenthos inhabiting the Antarctic sea floor, which is disturbed by iceberg scouring. We used models based on the empirical knowledge of succession to calculate the IDH curve for this system and to extrapolate the presently observable range of the IDH curve to higher and lower disturbance rates. Although the hump-shaped relationship has been found for a purely theoretical (extremely large) disturbance range, within the feasible disturbance range (assumed as realistic in the Antarctic region under climate change), the regional diversity of successional stages due to iceberg scouring strongly decreases with lower disturbance rates but levels off only slowly with higher disturbance rates. The reason is the unevenness in the lifetimes of the successional stages, in that early stages are short-lived whereas late stages are long-lived. With such unevenness, increasing disturbances support the early stages without jeopardizing the later ones. Additionally, we converted this regional diversity of stages to the regional diversity of taxa using a transformation formula based on empirical knowledge of the number and mean abundance of taxa in the particular stages. Our results suggest that a decrease in iceberg scouring due to climate change would be more detrimental to the diversity of the Antarctic megabenthos than an increase.     

Soft bottom species richness and diversity as a function of depth and iceberg scour in Arctic glacial Kongsfjorden (Svalbard)

Macrozoobenthic soft-sediment communities of central Arctic Kongsfjorden inhabiting six depth zones between 5 and 30 m were sampled using SCUBA-diving during June–August 2003 and analysed comparatively. About 63 taxa were found, nine of which had not been reported for Kongsfjorden and four for Svalbard. Suspension feeding or surface and sub-surface detritivorous polychaetes and deposit-feeding amphipods were dominant. Only 11 of the 63 taxa (45 species and additional 18 families not further identified) inhabited the complete depth range. Biomass ranged from 3.5 to 25.0 g ash free dry mass m⁻² and mean Shannon diversity (Log e) was 2.06. Similarity clustering from abundance and biomass data showed a significant difference between the shallow station (5 m) and the rest. The latter formed two sub-groups (10–20 and 25–30 m). Depth is irrevocably correlated with ice-scouring. Thus the differences in diversity together with the predicted iceberg scour intensity support the ‘intermediate disturbance hypothesis’ indicating that habitats impacted by moderate iceberg scouring enable higher diversity. In contrast, biotopes frequently affected only host pioneer communities, while mature, less diverse assemblages dominate depths of low impact.     

Small-scale spatial genetic structure in the Central African rainforest tree species Aucoumea klaineana: a stepwise approach to infer the impact of limited gene dispersal, population history and habitat fragmentation

Under the isolation-by-distance model, the strength of spatial genetic structure (SGS) depends on seed and pollen dispersal and genetic drift, which in turn depends on local demographic structure. SGS can also be influenced by historical events such as admixture of differentiated gene pools. We analysed the fine-scale SGS in six populations of a pioneer tree species endemic to Central Africa, Aucoumea klaineana. To infer the impacts of limited gene dispersal, population history and habitat fragmentation on isolation by distance, we followed a stepwise approach consisting of a Bayesian clustering method to detect differentiated gene pools followed by the analysis of kinship-distance curves. Interestingly, despite considerable variation in density, the five populations situated under continuous forest cover displayed very similar extent of SGS. This is likely due to an increase in dispersal distance with decreased tree density. Admixture between two gene pools was detected in one of these five populations creating a distinctive pattern of SGS. In the last population sampled in open habitat, the genetic diversity was in the same range as in the other populations despite a recent habitat fragmentation. This result may due to the increase of gene dispersal compensating the effect of the disturbance as suggested by the reduced extent of SGS estimated in this population. Thus, in A. klaineana, the balance between drift and dispersal may facilitate the maintenance of genetic diversity. Finally, from the strength of the SGS and population density, an indirect estimate of gene dispersal distances was obtained for one site: the quadratic mean parent-offspring distance, sigma(g), ranged between 210 m and 570 m.     

Variation of scavenger richness and abundance between sites of high and low iceberg scour frequency in Ryder Bay,west Antarctic Peninsula

Physical disturbance, particularly from iceberg scour, is a major structuring force in polar benthic communities at shelf depths. Scouring kills and damages benthic organisms providing food for the abundant scavenging fauna of coastal Antarctic waters. This trophic group is likely to be strongly affected by changes in iceberg scouring. A baited underwater camera system was used to examine the distribution of scavenging fauna in relation to the spatial variation in exposure to iceberg impacts experienced at different iceberg scouring conditions and depths within Ryder Bay. The results indicate that the relationships between depth and scavenger abundance and assemblage composition differed between high and low scour sites. Scavenger abundance increased with depth at high scour sites and fell with depth a low scour sites. There was also significant difference in community composition between sites within each scouring condition. Scavenger species richness also exhibited an increase with depth at most sites consistent with the established pattern of declining iceberg scouring frequency with depth. Shannon–Wiener diversity increased with depth but significantly more steeply in highly scoured sites. Our results suggest that depth and exposure to icebergs interact to shape the scavenger community. The significant differences within the high and low scour groups suggest that other factors remain to be investigated and that there is probably a nonlinear relationship between scouring intensity and the favourability of a site for scavengers.     

Clumped versus scattered: how does the spatial correlation of disturbance events affect biodiversity?

In this study, we systematically explore the effects of rate and spatial correlation (level of clumping) of disturbance events on a community of sessile species differing in their life history traits. A spatially explicit individual-based model shows that long-term coexistence is very sensitive to spatial correlation when the trade-off in life history traits includes differences in dispersal distances. Highest biodiversity emerges at highly correlated disturbances of intermediate rates. Diversity peaks shift to larger rates when clumping decreases. Scattered disturbances lead to competitive exclusion. Interestingly, we observed additional peaks in the diversity–disturbance curves at certain levels of clumping. Thus, subject to the differences in life history traits, particular combinations of disturbance rate and spatial correlation may enable subsets of species to coexist, which opens new possibilities for explaining diversity. Our results suggest that observation of high biodiversity under spatially correlated disturbances points to a competition–colonisation trade-off, which includes dispersal distances.     

Nonsignificant isolation by distance implies limited dispersal

Understanding the scale of dispersal is an important consideration in the conservation and management of many species. However, in species in which the high‐dispersal stage is characterized by tiny gametes or offspring, it may be difficult to estimate dispersal directly. This is the case for many marine species, whose pelagic larvae are dispersed by ocean currents by several days or weeks before beginning a benthic, more sedentary, adult stage. As consequence of the high‐dispersal larval stage, many marine species have low genetic structure on large spatial scales (Waples 1998 ; Hellberg 2007 ). Despite the high capacity for dispersal, some tagging studies have found that a surprising number of larvae recruit into the population they were released from (self‐recruitment). However, estimates of self‐recruitment are not informative about mean dispersal between subpopulations. To what extent are limited dispersal estimates from tagging studies compatible with high potential for dispersal and low genetic structure? In this issue, a study on five species of coral reef fish used isolation by distance (IBD) between individuals to estimate mean dispersal distances (Puebla et al. 2012 ). They found that mean dispersal was unexpectedly small (<50 km), given relatively low IBD slopes and long pelagic durations. This study demonstrates how low genetic structure is compatible with limited dispersal in marine species. A comprehensive understanding of dispersal in marine species will involve integrating methods that estimate dispersal over different spatial and temporal scales. Genomic data may increase power to resolve these issues but must be applied carefully to this question.     

Are Introduced Species Better Dispersers Than Native Species? A Global Comparative Study of Seed Dispersal Distance

We provide the first global test of the idea that introduced species have greater seed dispersal distances than do native species, using data for 51 introduced and 360 native species from the global literature. Counter to our expectations, there was no significant difference in mean or maximum dispersal distance between introduced and native species. Next, we asked whether differences in dispersal distance might have been obscured by differences in seed mass, plant height and dispersal syndrome, all traits that affect dispersal distance and which can differ between native and introduced species. When we included all three variables in the model, there was no clear difference in dispersal distance between introduced and native species. These results remained consistent when we performed analyses including a random effect for site. Analyses also showed that the lack of a significant difference in dispersal distance was not due to differences in biome, taxonomic composition, growth form, nitrogen fixation, our inclusion of non-invasive introduced species, or our exclusion of species with human-assisted dispersal. Thus, if introduced species do have higher spread rates, it seems likely that these are driven by differences in post-dispersal processes such as germination, seedling survival, and survival to reproduction.     

Epizoochorous seed dispersal in relation to seed availability – an experiment with a red fox dummy

Questions: Is the red fox a potential vector for epizoochorous seed dispersal? Can seed attachment and retention be predicted from plant and seed traits? Location: Grasslands in southern Norway. Methods: Epizoochorous seed attachment on the red fox was studied by walking a dummy fox through the vegetation and comparing seeds found on the dummy with the estimated seed availability in the vegetation. Seed retention, i.e. the ability of different seeds to stay on the fox, was estimated in a separate experiment. Seed attachment and retention were related to plant and seed traits using statistical models that account for heteroscedasticity and zero‐inflated data. Results: The majority of seeds attached to the fox originated from a few species, but also species without specific seed traits that are supposed to enhance epizoochory attached at least some seeds to the fox. The probability of seed attachment was positively related to plant height, bristle and hooked seed appendages, and negatively related to winged appendages, seed mass, and seed sphericity. Seed retention was positively related to the seed traits bristles, hooks and pappus. For several species, the results indicate a high potential for dispersal over long distances. Conclusions: In modern agricultural landscapes, large herbivores are often restricted in their mobility or are found at low densities, and other animal vectors may therefore be important for seed dispersal. In our study, a range of plant species were able to disperse by attaching seeds to, and having their seeds retained in, the fox fur some distance. We suggest that the red fox may be an important vector for epizoochorous seed dispersal in the agricultural landscape.     

Parentage analysis detects cryptic precapture dispersal in a philopatric rodent

Locating birthplaces using genetic parentage determination can increase the precision and accuracy with which animal dispersal patterns are established. We re-analyse patterns of movement away from the birthplace as a function of time, sex and population density for a sample of 303 banner-tailed kangaroo rats, Dipodomys spectabilis. We located birth sites using a combination of likelihood-based parentage analysis with live-trapping of mothers during the breeding season. The results demonstrate that natal-breeding site distances are density dependent in this species; in particular, both sexes emigrate earlier in the year, and females disperse farther than males, at low population densities. Banner-tailed kangaroo rats were chosen as a study system because live-trapping easily detects maternal and offspring locations; nevertheless, parentage analysis reveals that some offspring evade early detection and move substantial distances before their first capture. In a few cases, the approach even detects dispersal out of the natal 'deme' prior to first capture. Parentage analysis confirms the extreme philopatry of both sexes but indicates that prior estimates of median dispersal distance were too low. For D. spectabilis, more accurate location of individual birthplaces clarifies patterns of sex bias and density dependence in dispersal, and may resolve apparent discrepancies between direct and indirect estimates of dispersal distance. For species in which mothers can be more reliably trapped than juveniles, using offspring genotypes to locate parents is a novel way that genetic techniques can contribute to the analysis of animal dispersal.     

Dispersal traits may reflect dispersal distances,but dispersers may not connect populations demographically

Ecological traits that reflect movement potential are often used as proxies for measured dispersal distances. Whether such traits reflect actual dispersal is often untested. Such tests are important because maximum dispersal distances may not be achieved and many dispersal events may be unsuccessful (without reproduction). For insects, many habitat patches harbour ‘resident’ species that are present as larvae (sedentary) and adults (winged and dispersing), and ‘itinerant’ species present only as adults that have dispersed from elsewhere and fail to reproduce. We tested whether itinerancy patterns were temporally consistent, and whether itinerant and resident species differed in wing morphology, a strong correlate of flight capability. Over 3 years and at multiple locations in a 22 km stream length, we sampled larvae and adults of caddisflies in the genus Ecnomus to categorize species as residents or itinerants. Flight capacity was measured using wing size (length and area) and shape parameters (aspect ratio and the second moment of wing area). Three species of Ecnomus were residents and three species were itinerants, and patterns were consistent over 3 years. On average, itinerant species had larger wings, suggesting a greater capacity to fly long distances. Wing shape differed between species, but did not differ systematically between residents and itinerants. Wing morphology was associated with actual but not effective dispersal of some species of Ecnomus. Morphological traits may have weak explanatory power for hypotheses regarding the demographic connectedness of populations, unless accompanied by data demonstrating which dispersers contribute new individuals to populations.     

Dispersal may limit the occurrence of specialist wood decay fungi already at small spatial scales

We used species‐specific spore traps to measure airborne dispersal of the wood decay fungus Phlebia centrifuga (spore size 6.5–9 × 2.5–3 μm) up to 1000 m distance from a point source. We fitted two simple dispersal models, an empirical power law model and a semi‐mechanistic diffusion model to the data using the Bayesian approach. The diffusion model provided a better fit than the power law model which underestimated deposition at 3–55 m and overestimated deposition at longer and shorter distances. Model fit improved by allowing overdispersion, suggesting that spores are not dispersed independently but wind can transport spores in groups inside discrete air packages up to considerable distances. Using the fitted diffusion model and available information on the establishment rates of wood‐decay fungi, we examine the distance up to which colonisation from a single fruit body is likely to occur. We conclude that the diluting effect of distance and low establishment success make the occurrence of P. centrifuga dispersal limited possibly already at the distance of tens of metres and very probably at a few hundred metres from the nearest fruit body, despite the fact that under favourable conditions a high proportion of the spores can disperse considerably further. This conclusion is likely to hold generally for those fungal species that inhabit fragmented landscapes, have specialised resource and habitat requirements, and have similar spore size and other dispersal traits as P. centrifuga.     

Speciation below ground: Tempo and mode of diversification in a radiation of endogean ground beetles

Dispersal is a critical factor determining the spatial scale of speciation, which is constrained by the ecological characteristics and distribution of a species’ habitat and the intrinsic traits of species. Endogean taxa are strongly affected by the unique qualities of the below‐ground environment and its effect on dispersal, and contrasting reports indicate either high dispersal capabilities favoured by small body size and mediated by passive mechanisms, or low dispersal due to restricted movement and confinement inside the soil. We studied a species‐rich endogean ground beetle lineage, Typhlocharina, including three genera and more than 60 species, as a model for the evolutionary biology of dispersal and speciation in the deep soil . A time‐calibrated molecular phylogeny generated from >400 individuals was used to delimit candidate species, to study the accumulation of lineages through space and time by species–area–age relationships and to determine the geographical structure of the diversification using the relationship between phylogenetic and geographic distances across the phylogeny. Our results indicated a small spatial scale of speciation in Typhlocharina and low dispersal capacity combined with sporadic long distance, presumably passive dispersal events that fuelled the speciation process. Analysis of lineage growth within Typhlocharina revealed a richness plateau correlated with the range of distribution of lineages, suggesting a long‐term species richness equilibrium mediated by density dependence through limits of habitat availability. The interplay of area‐ and age‐dependent processes ruling the lineage diversification in Typhlocharina may serve as a general model for the evolution of high species diversity in endogean mesofauna.     

Strategy Space and the Disturbance Spectrum: A Life-History Model for Tree Species Coexistence

The disturbance spectrum consists of disturbance patterns differing in type, size, intensity, and frequency. It is proposed that tree life-history traits are adaptations to particular disturbance regimes. Four independent axes are proposed to define the dominant dimensions of tree strategy space: shade tolerance, tree height, capacity for vegetative reproduction, and seed dispersal distance. A fitness model was developed to elucidate interactions between the proposed life-history traits. The model shows how alternate life-history sets can coexist when disturbance patterns fluctuate in space and time. Variable disturbance regimes were shown, based on data and simulation results, to enhance species coexistence, as predicted. The strategy space model accurately predicts the number of common tree species for the eastern United States, boreal Canada, and southwestern pi?on-juniper woodlands. The model also provides an explanation for latitudinal gradients in tree species richness in North America and Europe. The proposed model predicts a relationship between disturbance characteristics and the species composition of a forest that allows for the coexistence of large numbers of species. The life-history traits of size, growth rate, life span, shade tolerance, age of reproduction, seed dispersal distance, and vegetative reproduction are all incorporated into the model.     

Dependence of sustainability on the configuration of marine reserves and larval dispersal distance

Marine reserves hold promise for maintaining biodiversity and sustainable fishery management, but studies supporting them have not addressed a crucial aspect of sustainability: the reduction in viability of populations with planktonic larvae dispersing along a coastal habitat with noncontiguous marine reserves. We show how sustainability depends on the fraction of natural larval settlement (FNLS) remaining after reserves are implemented, which in turn depends on reserve configuration and larval dispersal distance. Sustainability requires FNLS to be greater than an empir-ically determined minimum. Maintaining an adequate value for all species requires either a large, unlikely fraction (> 35%) of coastline in reserves, or reserves that are larger than the mean larval dispersal distance of the target species. FNLS is greater for species dispersing shorter distances, which implies reserves can lead to: (1) changes in community composition and (2) genetic selection for shorter dispersal distance. Dependence of sustainability on dispersal distance is a new source of uncertainty.     

Probability of successful larval dispersal declines fivefold over 1 km in a coral reef fish

A central question of marine ecology is, how far do larvae disperse? Coupled biophysical models predict that the probability of successful dispersal declines as a function of distance between populations. Estimates of genetic isolation-by-distance and self-recruitment provide indirect support for this prediction. Here, we conduct the first direct test of this prediction, using data from the well-studied system of clown anemonefish (Amphiprion percula) at Kimbe Island, in Papua New Guinea. Amphiprion percula live in small breeding groups that inhabit sea anemones. These groups can be thought of as populations within a metapopulation. We use the x- and y-coordinates of each anemone to determine the expected distribution of dispersal distances (the distribution of distances between each and every population in the metapopulation). We use parentage analyses to trace recruits back to parents and determine the observed distribution of dispersal distances. Then, we employ a logistic model to (i) compare the observed and expected dispersal distance distributions and (ii) determine the relationship between the probability of successful dispersal and the distance between populations. The observed and expected dispersal distance distributions are significantly different (p < 0.0001). Remarkably, the probability of successful dispersal between populations decreases fivefold over 1 km. This study provides a framework for quantitative investigations of larval dispersal that can be applied to other species. Further, the approach facilitates testing biological and physical hypotheses for the factors influencing larval dispersal in unison, which will advance our understanding of marine population connectivity.     

Mate-finding as an overlooked critical determinant of dispersal variation in sexually-reproducing animals

Dispersal is a critically important process in ecology, but robust predictive models of animal dispersal remain elusive. We identify a potentially ubiquitous component of variation in animal dispersal that has been largely overlooked until now: the influence of mate encounters on settlement probability. We use an individual-based model to simulate dispersal in sexually-reproducing organisms that follow a simple set of movement rules based on conspecific encounters, within an environment lacking spatial habitat heterogeneity. We show that dispersal distances vary dramatically with fluctuations in population density in such a model, even in the absence of variation in dispersive traits between individuals. In a simple random-walk model with promiscuous mating, dispersal distributions become increasingly 'fat-tailed' at low population densities due to the increasing scarcity of mates. Similar variation arises in models incorporating territoriality. In a model with polygynous mating, we show that patterns of sex-biased dispersal can even be reversed across a gradient of population density, despite underlying dispersal mechanisms remaining unchanged. We show that some widespread dispersal patterns found in nature (e.g. fat tailed distributions) can arise as a result of demographic variability in the absence of heterogeneity in dispersive traits across the population. This implies that models in which individual dispersal distances are considered to be fixed traits might be unrealistic, as dispersal distances vary widely under a single dispersal mechanism when settlement is influenced by mate encounters. Mechanistic models offer a promising means of advancing our understanding of dispersal in sexually-reproducing organisms.     

Colonization dynamics of a clonal pioneer plant on a glacier foreland inferred from spatially explicit and size-structured matrix models

The regional distribution of a plant species is a result of the dynamics of extinctions and colonizations in suitable habitats, especially in strongly fragmented landscapes. Here, we studied the role of spatial dynamics of the long-lived, clonal pioneer plant Geum reptans occurring on glacier forelands in the European Alps. We used demographic data from several years and sites in the Swiss Alps in combination with dispersal data to parametrize a matrix model for G. reptans to simulate extinctions, colonizations and spatial spread of established populations on glacial forelands. We used different scenarios with varying germination rates, wind and animal dispersal capabilities, and modes of spatial spread (seed-only vs clonal spread), resulting in population growth rates (λ) ranging from 1.04 to 1.20. Our results suggest that due to the low germination rate (~1%) and the very limited wind dispersal distances (99.8% of seeds are dispersed < 5 m), G. reptans has a low probability of establishing new populations and a very low spatial spread by seed dispersal alone. In contrast to the low rate of establishment, the persistence of established populations is high and even populations of only a few individuals have an extinction probability of less than 25% within 100 years. This high persistency is partly due to clonal reproduction via aboveground stolons. Clonal reproduction increases the population size and contributes considerably to the spatial spread of established populations. Our simulation results together with the known pattern of molecular diversity of G. reptans indicate that the occurrence of populations of this species in the Alps is unlikely to be a result of recent colonizations by long-distance dispersal, but rather a result of post-glacial colonizations by large migrating populations that were fragmented when glaciers retreated. Additionally, our simulations suggest that the currently observed high rates of glacial retreat might be too fast for pioneer plants, such as G. reptans, to keep up with the retreating ice and therefore might threaten existing populations.     

Propagule pressure and an invasion syndrome determine invasion success in a plant community model

The success of species invasions depends on multiple factors, including propagule pressure, disturbance, productivity, and the traits of native and non‐native species. While the importance of many of these determinants has already been investigated in relative isolation, they are rarely studied in combination. Here, we address this shortcoming by exploring the effect of the above‐listed factors on the success of invasions using an individual‐based mechanistic model. This approach enables us to explicitly control environmental factors (temperature as surrogate for productivity, disturbance, and propagule pressure) as well as to monitor whole‐community trait distributions of environmental adaptation, mass, and dispersal abilities. We simulated introductions of plant individuals to an oceanic island to assess which factors and species traits contribute to invasion success. We found that the most influential factors were higher propagule pressure and a particular set of traits. This invasion trait syndrome was characterized by a relative similarity in functional traits of invasive to native species, while invasive species had on average higher environmental adaptation, higher body mass, and increased dispersal distances, that is, had greater competitive and dispersive abilities. Our results highlight the importance in management practice of reducing the import of alien species, especially those that display this trait syndrome and come from similar habitats as those being managed.