Random Dispersal Research Articles

Coexistence and extinction in the Volterra-Lotka competition model with nonlocal dispersal

Coexistence and extinction for two species Volterra-Lotka competition systems withnonlocal dispersal are investigated in this paper. Sufficient conditions in terms of diffusion, reproduction, self-limitation, and competition rates are established for existence, uniqueness, and stability of coexistence states as well as for the extinction of one species. The focus is on environments with hostile surroundings. In this case, our results correspond to those for random dispersal under Dirichlet boundary conditions. Similar results hold for environments with non-flux boundary and for periodic environments, which correspond to those for random dispersal under Neumann boundary conditions and periodic boundary conditions, respectively.

A data-driven simulation of endozoochory by ungulates illustrates directed dispersal

Large herbivorous mammals act as vectors of dispersal for many forbs and grasses through ingestion and excretion of seeds (endozoochory). Attributes from the plant, animal and landscape need to be integrated in order to study patterns of dispersal in the field, for instance, with respect to the spatial deposition of single plant species in heterogeneous landscapes (‘seed shadows’). Drawing on a set of empirical data, we investigated the dispersal of 25 plant species by cattle in a coastal dune area by means of a spatially explicit simulation model. Model output for the number of dispersed seeds matched observations from field-collected dung pats well. We then applied the model to quantify the seed shadow among the landscape's main vegetation types, focusing on whether the transfer of plants is directed towards suitable habitat. We herewith examined how changing habitat availability in the face of vegetation succession and management affects deposition patterns. A simple wind dispersal model was included for comparison. Simulation showed endozoochorous deposition to be most likely in grassland, but not when this habitat's surface area was low. Since the probability to end up in grassland was higher than expected from random dispersal, grassland plants were always provided with directed dispersal (but not always as strong). Our model illustrates the potential of grazers to contribute in directed seed transfer, apart from their role in long-distance dispersal.

Variation in the degree of specialization can maintain local diversity in model communities

We hypothesize that the continuum between generalist and specialist adaptations is an important general trade-off axis in the maintenance of local diversity, and we explore this idea with a simple model in which there are patch types to which species arrive as propagules and compete. Each patch type is defined by a competitive ranking of all species. A highly specialist species is the top competitor in one patch type but has a relatively low average ranking across different patch types, while a generalist species has a high average rank across patch types but is not the top competitor in any patch type. We use random dispersal and vary the fecundity of all species together to vary total propagule density and therefore recruitment limitation and density-dependent mortality. When fecundity is very high, each patch becomes occupied by its specialist species and generalists go extinct, so the number of species at equilibrium is equal to the number of patch types. If fecundity is very low, generalists dominate and specialists go extinct. There is a range of fecundity levels in which specialists, generalists, and intermediates coexist, and the number of species is substantially greater than the number of patch types. While coexistence of specialists and generalists has been considered a problem in evolutionary ecology, our results suggest to the contrary that this trade-off contributes to the maintenance of local diversity.

Accounting for spatial autocorrelation in null models of tree species association

A commonly used null model for species association among forest trees is a well‐mixed community (WMC). A WMC represents a non‐spatial, or spatially implicit, model, in which species form nearest‐neighbor pairs at a rate equal to the product of their community proportions. WMC models assume that the outcome of random dispersal and demographic processes is complete spatial randomness (CSR) in the species’ spatial distributions. Yet, stochastic dispersal processes often lead to spatial autocorrelation (SAC) in tree species densities, giving rise to clustering, segregation, and other nonrandom patterns. Although methods exist to account for SAC in spatially‐explicit models, its impact on non‐spatial models often remains unaccounted for. To investigate the potential for SAC to bias tests based upon non‐spatial models, we developed a spatially‐heterogeneous (SH) modelling approach that incorporates measured levels of SAC. Using the mapped locations of individuals in a tropical tree community, we tested the hypothesis that the identity of nearest‐neighbors represents a random draw from neighborhood species pools. Correlograms of Moran's I confirmed that, for 50 of 51 dominant species, stem density was significantly autocorrelated over distances ranging from 50 to 200 m. The observed patterns of SAC were consistent with dispersal limitation, with most species occurring in distinct patches. For nearly all of the 106 species in the community, the frequency of pairwise association was statistically indistinguishable from that projected by the null models. However, model comparisons revealed that non‐spatial models more strongly underestimated observed species‐pair frequencies, particularly for conspecific pairs. Overall, the CSR models projected more significant facilitative interactions than did SH models, yielding a more liberal test of niche differences. Our results underscore the importance of accounting for stochastic spatial processes in tests of association, regardless of whether spatial or non‐spatial models are employed.

Not across the North Pole: Plant migration in the Arctic

• The vascular plant flora of 66 arctic islands was studied to determine whether the islands have been occupied by random long-distance dispersal (LDD) or in a highly structured northward migration pattern via intervening islands as stepping-stones. • A maximum parsimonious migration model minimizing dispersal distances of 1256 vascular plant taxa was calculated in the framework of network analysis. • Plant dispersal is not stochastic in the Arctic at the global scale. Inferred mean dispersal distances of the plants occurring on arctic islands are c. 580 km (median 460 km). A LDD across the North Pole could not be inferred in the model and species may be recruited mainly from the nearest mainland or islands. At smaller scales, among adjacent islands, dispersal of vascular plants may be incomplete. Arctic islands do not yet appearto be saturated with species. • The results suggest that changes in biodiversity in Arctic islands can be more easily predicted at the global scale than at the local scale. Because islands are not yet saturated with species, new colonizations may not necessarily be linked to climate change.

Empirical evaluation of directed dispersal and density‐dependent effects across successive recruitment phases

Summary1. The directed‐dispersal (DrD) hypothesis constitutes one of the main explanations for the adaptive value of seed dispersal in spatially heterogeneous environments. Traditionally, the DrD hypothesis asserts non‐random arrival to specific sites where establishment conditions are independently favourable. Yet, enhanced arrival might negatively affect the establishment in otherwise favourable sites through enhanced density‐dependent mortality (DDM). Since both density effects and habitat suitability might differ among establishment phases, assessment of habitat suitability should encompass the entire establishment process.2. Here, we test the DrD hypothesis against a null model of random dispersal (RD) in an arid ecosystem where fleshy‐fruited perennial Ochradenus baccatus shrubs are dispersed by two resident passerines. Acacia trees were predicted to present the favourable habitat in our system through the nurse‐plant effect facilitating O. baccatus early establishment.3. We determined expected seed arrival by monitoring the dispersers’ time allocation among the different habitats and assessed habitat quality for different establishment phases: seed‐phase survival (post‐dispersal survival and germination) was determined by line transects, while early seedling survival was estimated by monitoring seedlings throughout the first dry season. Additionally, subsequent recruitment success was assessed by comparing the distribution of adult shrubs to that expected from the observed germination patterns.4. When considering the expected seed arrival and early establishment (seed‐phase and early seedling survival), the DrD hypothesis is supported: birds preferentially deposited seeds in habitats where conditions are favourable (e.g. under Acacia trees). Yet, considering recruitment success during later phases, the dispersal service provided by the birds was less beneficial than RD.5. Effective density was indexed using the zone‐of‐influence approach in which densities are corrected by phase‐specific sizes. We suggest that intense seed deposition into favourable habitats has led to strong DDM (presumably through competition) in those habitats during late establishment phases and was responsible, at least partially, for changes in habitat suitability across the recruitment phases.6. Synthesis. Our results show that assessment of the DrD hypothesis depends on the context of the study (in terms of recruitment phases) since sites of enhanced seed arrival and high seed‐to‐seedling survival can ultimately prove unfavourable to the plant if density‐dependent or other effects might increase mortality in these sites during later recruitment phases.

Evaluating environmental and geometrical constraints on endemic vertebrates of the semiarid Caatinga (Brazil)

Most hypotheses explaining species diversity are related to the current and/or past environment, but explanations that rely on geometrical constraints, such as the mid-domain effect, have been recently proposed. Under this type of model, more species are expected in the centre of a domain due to the extent of occurrence of the species being constrained by the domain's geometry. Here we evaluated the proportion of the explanatory power of space, the current environment, historical vicariance events and geometric constraints on the richness of the Caatinga's terrestrial endemic vertebrates using a partial regression approach. The current environment had the highest proportion of explanatory power for the variance of the Caatinga's endemic species, in its shared effect, explaining approximately 47%. On the other hand the geometric constraints model (GCM) had the highest proportion in its single explanatory power with 15% of explanatory power. The amount of explanation by GCM changes when analyzing richness of endemics calculated from species with different range sizes. Despite the fact that GCM is the main driver of the Caatinga's endemic species, in its single effect, this does not necessarily indicate that geometric constraints and random dispersal within species ranges drive richness. The GCM model underestimates richness in the centre of the Caatinga, suggesting that other ecological and evolutionary processes may help explain the overall pattern of decreasing richness towards the borders of the biome.

Determinism versus chance in canopy gap herbaceous species assemblages in temperate Abies– Betula forests

It is not known if the species composition of herbaceous plant assemblies within tree-fall gaps is determined stochastically or whether it follows species-specific and environmentally-determined patterns. We applied three methods, fitting species abundance distribution models, comparing community similarities, and testing species–environmental variable association, to evaluate the relative importance of chance and deterministic rules in controlling herbaceous composition within canopy gaps in the Abies– Betula forests of Mt. Taibai, Central China. Herbaceous species abundance within canopy gaps was well fit by a neutral model, showing that relative abundance was qualitatively consistent with stochastic processes. Although species composition in gaps significantly differed from closed canopy sites (ANOSIM, R = 0.509, p = 0.001), there were no significant differences among gaps of different age groups (ANOSIM, R = 0.035, p = 0.191) or gaps of different size groups (ANOSIM, R = 0.089, p = 0.057). Similarly, gaps of similar age and size did not show significantly higher similarity ( χ 2 = 2.30, df = 3, p = 0.513) in species composition than gaps of distinct age or size. Moreover, there were only eight species, including two light-demanding species, confined to larger gaps among the 69 species, and only an additional seven species found more commonly in larger gaps, whereas most herbaceous species were gap size generalists. Canonical correspondence analysis and random permutation tests suggested that only 27% of species with abundance ⩾5 were associated with environmental variables in gaps. In summary, the species composition in gaps was not constrained significantly by gap traits; rather species were distributed stochastically, likely through by random dispersal and recruitment limitation of species from the surrounding available species pool. Measures that introduce gap disturbance, such as selective harvesting, are still recommended when the maintenance of total biodiversity in forests is a concern.

Landscape connectivity and seed dispersal characteristics inform the best management strategy for exotic plants

Exotic plant invasions have triggered environmental and economic problems throughout the world. Our ability to manage these invasions is hindered by the difficulty of predicting spread in fragmented landscapes. Because the spatial pattern of invasions depends on the dispersal characteristics of the invasive species and the configuration of suitable habitat within the landscape, a universal management strategy is unlikely to succeed for any particular species. We suggest that the most effective management strategy may be an adaptive one that shifts from local control to landscape management depending on the specific invader and landscape. In particular, we addressed the question of where management activities should be focused to minimize spread of the invading species. By simulating an invasion across a real landscape (Antietam National Battlefield in Maryland, USA), we examined the importance of patch size and connectivity to management success. We found that the best management strategy depended on the dispersal characteristics of the exotic species. Species with a high probability of random long-distance dispersal were best managed by focusing on the largest patches, while species with a lower probability of random long-distance dispersal were best managed by considering landscape configuration and connectivity of the patches. Connectivity metrics from network analysis were useful for identifying the most effective places to focus management efforts. These results provide insight into invasion patterns of various species and suggest a general rule for managers in National Parks and other places where invasive species are a concern.

Measuring biodiversity to explain community assembly: a unified approach

One of the oldest challenges in ecology is to understand the processes that underpin the composition of communities. Historically, an obvious way in which to describe community compositions has been diversity in terms of the number and abundances of species. However, the failure to reject contradictory models has led to communities now being characterized by trait and phylogenetic diversities. Our objective here is to demonstrate how species, trait and phylogenetic diversity can be combined together from large to local spatial scales to reveal the historical, deterministic and stochastic processes that impact the compositions of local communities. Research in this area has recently been advanced by the development of mathematical measures that incorporate trait dissimilarities and phylogenetic relatedness between species. However, measures of trait diversity have been developed independently of phylogenetic measures and conversely most of the phylogenetic diversity measures have been developed independently of trait diversity measures. This has led to semantic confusions particularly when classical ecological and evolutionary approaches are integrated so closely together. Consequently, we propose a unified semantic framework and demonstrate the importance of the links among species, phylogenetic and trait diversity indices. Furthermore, species, trait and phylogenetic diversity indices differ in the ways they can be used across different spatial scales. The connections between large-scale, regional and local processes allow the consideration of historical factors in addition to local ecological deterministic or stochastic processes. Phylogenetic and trait diversity have been used in large-scale analyses to determine how historical and/or environmental factors affect both the formation of species assemblages and patterns in species richness across latitude or elevation gradients. Both phylogenetic and trait diversity have been used at different spatial scales to identify the relative impacts of ecological deterministic processes such as environmental filtering and limiting similarity from alternative processes such as random speciation and extinction, random dispersal and ecological drift. Measures of phylogenetic diversity combine phenotypic and genetic diversity and have the potential to reveal both the ecological and historical factors that impact local communities. Consequently, we demonstrate that, when used in a comparative way, species, trait and phylogenetic structures have the potential to reveal essential details that might act simultaneously in the assembly of species communities. We highlight potential directions for future research. These might include how variation in trait and phylogenetic diversity alters with spatial distances, the role of trait and phylogenetic diversity in global-scale gradients, the connections between traits and phylogeny, the importance of trait rarity and independent evolutionary history in community assembly, the loss of trait and phylogenetic diversity due to human impacts, and the mathematical developments of biodiversity indices including within-species variations.

Functional uniqueness of a small carnivore as seed dispersal agents: a case study of the common palm civets in the Tabin Wildlife Reserve, Sabah, Malaysia

Many carnivorous mammals consume fruits and disperse the intact seeds to specific sites. Few studies have attempted to quantify this seed dispersal or evaluate its effectiveness, despite its potential importance and functional uniqueness. In the study reported here, we found that a frugivorous carnivore, the common palm civet (Paradoxurus hermaphroditus), generated seed shadows that are distinct from those of the sympatric frugivore, the pig-tailed macaque (Macaca nemestrina), and played a unique and important role in the regeneration of Leea aculeata (Leeaceae). We found that macaques dispersed the seeds randomly, while civets dispersed them non-randomly to sites such as the banks of small rivers, rain-flow paths, abandoned trails, and treefall gaps, which are characterized by low stem density and canopy cover. Seeds of L. aculeata that were dispersed by civets to the banks of rivers and gaps had significantly higher survival and growth rates than those dispersed to rain-flow paths or abandoned trails. Seeds dispersed by macaques or to random locations also had low survival. Although the effects of the civets on seed fate were not straightforward, compared with macaques and random dispersal, civets significantly enhanced the survival and growth of L. aculeata seeds after 1 year. These results indicate that non-random dispersal by civets is important for the persistence of L. aculeata. Civets may disperse other plant species and thus could have profound effects on forest dynamics.

Modeling the principle physical parameters of graphite carbon foam

Graphite carbon foam, a mesophase, pitch-based material, portrays highly ordered topology structures which exhibit superior mechanical and thermal properties. Typical graphite carbon foam with dimensions 5cm3, can have a surface area greater than 11m2, making it an excellent candidate for heat transfer applications. Accurate three dimensional modeling of carbon foams is necessary to study and predict their properties in simulation. This paper describes a computer algorithm for modeling POCO Foam® and similar carbon foams. The algorithm, written in MATLAB, captures the principle physical parameters of the carbon foam including bubble and pore diameter ranges and overall foam void percentage while retaining the random dispersal of spherical bubbles found in manufactured foams.

Deep evolutionary roots of strepsirrhine primate labyrinthine morphology

The cavity system of the inner ear of mammals is a complex three-dimensional structure that houses the organs of equilibrium and hearing. Morphological variation of the inner ear across mammals reflects differences in locomotor behaviour and hearing performance, and the good preservation of this structure in many fossil specimens permits analogous inferences. However, it is less well known to what extent the morphology of the bony labyrinth conveys information about the evolutionary history of primate taxa. We studied this question in strepsirrhine primates with the aim to assess the potential and limitations of using the inner ear as a phylogenetic marker. Geometric morphometric analysis showed that the labyrinthine morphology of extant strepsirrhines contains a mixed locomotor, allometric and phylogenetic signal. Discriminant analysis at the family level confirmed that labyrinthine shape is a good taxonomic marker. Our results support the hypothesis that evolutionary change in labyrinthine morphology is adequately described with a random walk model, i.e. random phenotypic dispersal in morphospace. Under this hypothesis, average shapes calculated for each node of the phylogenetic tree give an estimate of inner ear shapes of the respective last common ancestors (LCAs), and this information can be used to infer character state polarity. The labyrinthine morphology of the fossil Adapinae is close to the inferred basal morphology of the strepsirrhines. The inner ear of Daubentonia, one of the most derived extant strepsirrhines, is autapomorphic in many respects, but also presents unique similarities with adapine labyrinths.

Genetic diversity in three invasive clonal aquatic species in New Zealand.

BackgroundElodea canadensis, Egeria densa and Lagarosiphon major are dioecious clonal species which are invasive in New Zealand and other regions. Unlike many other invasive species, the genetic variation in New Zealand is very limited. Clonal reproduction is often considered an evolutionary dead end, even though a certain amount of genetic divergence may arise due to somatic mutations. The successful growth and establishment of invasive clonal species may be explained not by adaptability but by pre-existing ecological traits that prove advantageous in the new environment. We studied the genetic diversity and population structure in the North Island of New Zealand using AFLPs and related the findings to the number of introductions and the evolution that has occurred in the introduced area.ResultsLow levels of genetic diversity were found in all three species and appeared to be due to highly homogeneous founding gene pools. Elodea canadensis was introduced in 1868, and its populations showed more genetic structure than those of the more recently introduced of E. densa (1946) and L. major (1950). Elodea canadensis and L. major, however, had similar phylogeographic patterns, in spite of the difference in time since introduction.ConclusionsThe presence of a certain level of geographically correlated genetic structure in the absence of sexual reproduction, and in spite of random human dispersal of vegetative propagules, can be reasonably attributed to post-dispersal somatic mutations. Direct evidence of such evolutionary events is, however, still insufficient.

Effect of non‐random dispersal strategies on spatial coexistence mechanisms

1. Random dispersal leads to spatial coexistence via two mechanisms (emigration-mediated and source-sink), both of which involve the movement of organisms from areas of higher to lower fitness. What is not known is whether such coexistence would occur if organisms dispersed non-randomly, using cues such as density and habitat quality to gauge fitness differences between habitats. Here, I conduct a comparative analysis of random and non-random dispersal strategies in a foodweb with a basal resource, top predator, and two intermediate consumers that exhibit a trade-off between competitive ability and predator susceptibility. 2. I find a striking contrast between density- and habitat-dependent dispersal in their effects on spatial coexistence. Dispersal in response to competitor and predator density facilitates coexistence while dispersal in response to habitat quality (resource productivity and predator pressure) inhibits it. Moreover, density-dependent dispersal changes species' distribution patterns from interspecific segregation to interspecific aggregation, while habitat-dependent dispersal preserves the interspecific segregation observed in the absence of dispersal. Under density-dependent dispersal, widespread spatial coexistence results in an overall decline in the abundance of the inferior competitor that is less susceptible to predation and an overall increase in the abundance of the superior competitor that is more susceptible to predation. Under habitat-dependent dispersal, restricted spatial coexistence results in species' abundances being essentially unchanged from those observed in the absence of dispersal. 3. A key outcome is that when the superior competitor moves in the direction of increasing fitness but the inferior competitor does not, spatial coexistence is possible in both resource-poor and resource-rich habitats. However, when the inferior competitor moves in the direction of increasing fitness but the superior competitor does not, spatial coexistence is precluded in resource-poor habitats and greatly reduced in resource-rich habitats. This suggests that species-specific differences may play an important role in driving spatial coexistence patterns. 4. The comparative framework yields predictions that can be tested with experiments that manipulate the relative mobilities of interacting species, or observational data on relative abundances and distribution patterns.

Random dispersal vs. non-local dispersal

Random dispersal is essentially a localbehavior which describes the movement of organisms betweenadjacent spatial locations. However, the movements andinteractions of some organisms can occur between non-adjacentspatial locations. To address the question about which dispersalstrategy can convey some competitive advantage, we consider amathematical model consisting of one reaction-diffusion equationand one integro-differential equation, in which two competingspecies have the same population dynamics but different dispersalstrategies: the movement of one species is purely by random walkwhile the other species adopts a non-local dispersal strategy.For spatially periodic and heterogeneous environments we showthat (i) for fixed random dispersal rate,if the nonlocal dispersal distance is sufficiently small, thenthe non-local disperser can invade the random disperser but not vice versa;(ii) for fixed nonlocal dispersal distance,if the random dispersal rate becomes sufficiently small, thenthe random disperser can invade the nonlocal disperser but notvice versa.These results suggestthat for spatially periodic heterogeneous environments,the competitive advantage may belong to the species with much lowereffective rate of dispersal. This is in agreement with previous resultsfor the evolution of random dispersal [9, 13] that the slower disperser has an advantage. Nevertheless, if random dispersal strategy with either zero Dirichlet or zero Neumann boundary condition is compared with non-local dispersal strategy with hostilesurroundings, the species with much lower effective rate of dispersal may not have the competitive advantage. Numerical results will be presented to shed light on the global dynamics of the system for general values of non-local interaction distance and also to point to future research directions.

Modelling single nucleotide effects inphosphoglucose isomeraseon dispersal in the Glanville fritillary butterfly: coupling of ecological and evolutionary dynamics

Dispersal comprises a complex life-history syndrome that influences the demographic dynamics of especially those species that live in fragmented landscapes, the structure of which may in turn be expected to impose selection on dispersal. We have constructed an individual-based evolutionary sexual model of dispersal for species occurring as metapopulations in habitat patch networks. The model assumes correlated random walk dispersal with edge-mediated behaviour (habitat selection) and spatially correlated stochastic local dynamics. The model is parametrized with extensive data for the Glanville fritillary butterfly. Based on empirical results for a single nucleotide polymorphism (SNP) in the phosphoglucose isomerase (Pgi) gene, we assume that dispersal rate in the landscape matrix, fecundity and survival are affected by a locus with two alleles, A and C, individuals with the C allele being more mobile. The model was successfully tested with two independent empirical datasets on spatial variation in Pgi allele frequency. First, at the level of local populations, the frequency of the C allele is the highest in newly established isolated populations and the lowest in old isolated populations. Second, at the level of sub-networks with dissimilar numbers and connectivities of patches, the frequency of C increases with decreasing network size and hence with decreasing average metapopulation size. The frequency of C is the highest in landscapes where local extinction risk is high and where there are abundant opportunities to establish new populations. Our results indicate that the strength of the coupling of the ecological and evolutionary dynamics depends on the spatial scale and is asymmetric, demographic dynamics having a greater immediate impact on genetic dynamics than vice versa.

The Evolution of Conditional Dispersal Strategies in Spatially Heterogeneous Habitats

To understand the evolution of dispersal, we study a Lotka-Volterra reaction-diffusion-advection model for two competing species in a heterogeneous environment. The two species are assumed to be identical except for their dispersal strategies: both species disperse by random diffusion and advection along environmental gradients, but with slightly different random dispersal or advection rates. Two new phenomena are found for one-dimensional habitats and monotone intrinsic growth rates: (i) If both species disperse only by random diffusion, i.e., no advection, it was well known that the slower diffuser always wins. We show that if both species have the same advection rate which is suitably large, the faster dispersal will evolve; (ii) If both species have the same random dispersal rate, it was known that the species with a little advection along the resource gradient always wins, provided that the other species is a pure random disperser and the habitat is convex. We show that if both species have the same random dispersal rate and both also have suitably large advection rates, the species with a little smaller advection rate always wins. Implications of these results for the habitat choices of species will be discussed. Some future directions and open problems will be addressed.

Recurring cluster and operon assembly for Phenylacetate degradation genes

BackgroundA large number of theories have been advanced to explain why genes involved in the same biochemical processes are often co-located in genomes. Most of these theories have been dismissed because empirical data do not match the expectations of the models. In this work we test the hypothesis that cluster formation is most likely due to a selective pressure to gradually co-localise protein products and that operon formation is not an inevitable conclusion of the process.ResultsWe have selected an exemplar well-characterised biochemical pathway, the phenylacetate degradation pathway, and we show that its complex history is only compatible with a model where a selective advantage accrues from moving genes closer together. This selective pressure is likely to be reasonably weak and only twice in our dataset of 102 genomes do we see independent formation of a complete cluster containing all the catabolic genes in the pathway. Additionally, de novo clustering of genes clearly occurs repeatedly, even though recombination should result in the random dispersal of such genes in their respective genomes. Interspecies gene transfer has frequently replaced in situ copies of genes resulting in clusters that have similar content but very different evolutionary histories.ConclusionOur model for cluster formation in prokaryotes, therefore, consists of a two-stage selection process. The first stage is selection to move genes closer together, either because of macromolecular crowding, chromatin relaxation or transcriptional regulation pressure. This proximity opportunity sets up a separate selection for co-transcription.

Fine‐scale genetic structure of a long‐lived reptile reflects recent habitat modification

Anthropogenic habitat fragmentation--ubiquitous in modern ecosystems--has strong impacts on gene flow and genetic population structure. Reptiles may be particularly susceptible to the effects of fragmentation because of their extreme sensitivity to environmental conditions and limited dispersal. We investigate fine-scale spatial genetic structure, individual relatedness, and sex-biased dispersal in a large population of a long-lived reptile (tuatara, Sphenodon punctatus) on a recently fragmented island. We genotyped individuals from remnant forest, regenerating forest, and grassland pasture sites at seven microsatellite loci and found significant genetic structuring (R(ST)=0.012) across small distances (<500 m). Isolation by distance was not evident, but rather, genetic distance was weakly correlated with habitat similarity. Only individuals in forest fragments were correctly assignable to their site of origin, and individual pairwise relatedness in one fragment was significantly higher than expected. We did not detect sex-biased dispersal, but natural dispersal patterns may be confounded by fragmentation. Assignment tests showed that reforestation appears to have provided refuges for tuatara from disturbed areas. Our results suggest that fine-scale genetic structuring is driven by recent habitat modification and compounded by the sedentary lifestyle of these long-lived reptiles. Extreme longevity, large population size, simple social structure and random dispersal are not strong enough to counteract the genetic structure caused by a sedentary lifestyle. We suspect that fine-scale spatial genetic structuring could occur in any sedentary species with limited dispersal, making them more susceptible to the effects of fragmentation.