Long distance dispersal and the fate of a gene from the colonization front

There is an increasing recognition that long distance dispersal (LDD) plays a key role in establishing spatial genetic structure during colonization. Recent works, focused on short distance dispersal, demonstrated that a neutral mutation arising at the colonization front can either ‘surf’ with the wave front and reach high frequencies or stay near its place of origin at low frequencies. Here, we examine how LDD, and more generally the shape of the dispersal kernel, modifies this phenomenon and how the width of the colonization corridor affects the fate of the mutation. We demonstrate that when LDD events are more frequent, the ‘surfing phenomenon’ is less frequent, probably because any alleles can get far ahead from the colonization front and preclude the invasion by others alleles, thus leading to an attenuation of the diversity loss. We also demonstrate that the width of the colonization corridor influences the fate of the mutation, wide spaces decreasing the probability of invasion. Overall, the genetic structure of diversity resulted not only from LDD but also particularly from the shape of the dispersal kernel.     

A first step towards inferring levels of long-distance dispersal during past expansions

Improving the realism of spatially explicit demographic models is important for better inferring the history of past populations and for understanding the genetic bases of adaptation and speciation. One particular type of demographic event to take into account is long-distance dispersal (LDD). The goals of this study are to explore the impact of various levels of LDD on genetic diversity and to show to what extent LDD levels can be correctly inferred from multilocus data sets using an approximate Bayesian computation approach. We therefore incorporated LDD into a 2D stepping stone forward simulation framework coupled to a coalescent backward simulation step to generate genetic diversity at 100 microsatellite markers under various demographic conditions relevant to recent human evolution. Our results confirm that LDD considerably increases genetic diversity within demes and decreases levels of diversity between demes. By controlling the spatial occurrence of LDD, it appears that LDD events occurring during a phase of range expansion into new territories are more important in maintaining genetic diversity than those occurring in the wake of the expansion or when colonization is over. We also show that it is possible to infer whether LDD has occurred during a range expansion, but our results suggest that one can only approximately estimate the extent of LDD based on genetic summary statistics.     

The importance of long-distance dispersal in biodiversity conservation

Dispersal is universally considered important for biodiversity conservation. However, the significance of long‐ as opposed to short‐distance dispersal is insufficiently recognized in the conservation context. Long‐distance dispersal (LDD) events, although typically rare, are crucial to population spread and to maintenance of genetic connectivity. The main threats to global biodiversity involve excessive LDD of elements alien to ecosystems and insufficient dispersal of native species, for example, because of habitat fragmentation. In this paper, we attempt to bridge the gap in the treatment of LDD by reviewing the conservation issues for which LDD is most important. We then demonstrate how taking LDD into consideration can improve conservation management decisions.     

The population genetic structure of clonal organisms generated by exponentially bounded and fat-tailed dispersal

Long-distance dispersal (LDD) plays an important role in many population processes like colonization, range expansion, and epidemics. LDD of small particles like fungal spores is often a result of turbulent wind dispersal and is best described by functions with power-law behavior in the tails ("fat tailed"). The influence of fat-tailed LDD on population genetic structure is reported in this article. In computer simulations, the population structure generated by power-law dispersal with exponents in the range of -2 to -1, in distinct contrast to that generated by exponential dispersal, has a fractal structure. As the power-law exponent becomes smaller, the distribution of individual genotypes becomes more self-similar at different scales. Common statistics like GST are not well suited to summarizing differences between the population genetic structures. Instead, fractal and self-similarity statistics demonstrated differences in structure arising from fat-tailed and exponential dispersal. When dispersal is fat tailed, a log-log plot of the Simpson index against distance between subpopulations has an approximately constant gradient over a large range of spatial scales. The fractal dimension D2 is linearly inversely related to the power-law exponent, with a slope of approximately -2. In a large simulation arena, fat-tailed LDD allows colonization of the entire space by all genotypes whereas exponentially bounded dispersal eventually confines all descendants of a single clonal lineage to a relatively small area.     

Impacts of Restored Patch Density and Distance from Natural Forests on Colonization Success

Abstract The reduction and fragmentation of forest habitats is expected to have profound effects on plant species diversity as a consequence of the decreased area and increased isolation of the remnant patches. To stop the ongoing process of forest fragmentation, much attention has been given recently to the restoration of forest habitat. The present study investigates restoration possibilities of recently established patches with respect to their geographical isolation. Because seed dispersal events over 100 m are considered to be of long distance, a threshold value of 100 m between recent and old woodland was chosen to define isolation. Total species richness, individual patch species richness, frequency distributions in species occurrences, and patch occupancy patterns of individual species were significantly different among isolated and nonisolated stands. In the short term no high species richness is to be expected in isolated stands. Establishing new forests adjacent to existing woodland ensures higher survival probabilities of existing populations. In the long term, however, the importance of long‐distance seed dispersal should not be underestimated because most species showed occasional long‐distance seed dispersal. A clear distinction should be made between populations colonizing adjacent patches and patches isolated from old woodland. The colonization of isolated stands may have important effects on the dynamics and diversity of forest networks, and more attention should be directed toward the genetic traits and viability of founding populations in isolated stands.     

Genetic discontinuities and disequilibria in recently established populations of the plant pathogenic fungus Mycosphaerella fijiensis

Dispersal processes of fungal plant pathogens can be inferred from analysis of spatial genetic structures resulting from recent range expansion. The relative importance of long‐distance dispersal (LDD) events vs. gradual dispersal in shaping population structures depends on the geographical scale considered. The fungus Mycosphaerella fijiensis, pathogenic on banana, is an example of a recent worldwide epidemic. Founder effects in this species were detected at both global and continental scale, suggesting stochastic spread of the disease through LDD events. In this study, we analysed the structure of M. fijiensis populations in two recently (∼1979–1980) colonized areas in Costa Rica and Cameroon. Isolates collected in 10–15 sites distributed along a ∼250‐ to 300‐ km‐long transect in each country were analysed using 19 microsatellite markers. We detected low‐to‐moderate genetic differentiation among populations in both countries and isolation by distance in Cameroon. Combined with historical data, these observations suggest continuous range expansion at the scale of banana‐production area through gradual dispersal of spores. However, both countries displayed specific additional signatures of colonization: a sharp discontinuity in gene frequencies was observed along the Cameroon transect, while the Costa Rican populations seemed not yet to have reached genetic equilibrium. These differences in the genetic characteristics of M. fijiensis populations in two recently colonized areas are discussed in the light of historical data on disease spread and ecological data on landscape features.     

Lichen colonization patterns show minor effects of dispersal distance at landscape scale

The role of dispersal in controlling the distribution of species at landscape scale (10²–10⁴ m) is still a matter of dispute. Here, we use the early colonization pattern of 23 epiphytic lichen species in a former tree‐less heathland landscape (170 km²) to test three hypotheses on how a landscape is colonized: A) mainly by long‐distance dispersal (LDD), B) by rare LDD events followed by limited local dispersal, and C) mainly by limited dispersal, resulting in a colonization front. The study system consisted of a chronosequence of 94 habitat patches constituting 0.4% of the landscape area, with a minimum inter‐site distance of 0.2 km. We used generalized linear mixed models with Bayesian inference to test predictions from the hypotheses. When age of sites and habitat area were accounted for, additional effects of geographical position of sites (distance from old sites, distance‐dependent relative propagule pressure, and distance from border of study area) on the probability of colonization by lichen species were small. Furthermore, species richness of sites did not depend on geographical position, either. Our results support a colonization process mainly governed by LDD at landscape scale, and that local stepwise colonization was not important. We argue that passively dispersed species with numerous small propagules tend to exhibit patchy populations with extensive dispersal at the landscape scale, rather than behaving like classical metapopulations.     

Range expansion and retraction along a moving contact zone has no effect on the genetic diversity of two passerine birds

Disentangling the factors shaping species distributions remains a central goal in biogeography, ecology and evolutionary biology. The extrinsic pressures that may facilitate range shifts, such as climatic factors or biotic interactions are well known. However, in contrast, the possible intrinsic factors are manifold and hard to generalize across taxa. Recently, several theoretical studies have investigated the consequences of moving range borders on genetic diversity. However, empirical studies that support or refute these theoretical predictions are scarce. Moving contact zones between parapatric sister species are suitable models to test these hypotheses. Changes in genetic diversity can be tested simultaneously along the expanding and receding edges of two species of the contact zone while accounting for intra‐specific effects (e.g. introgression). The two Old World warblers Hippolais polyglotta and H. icterina form a narrow moving contact zone, where interspecific interactions are suspected to be the main factor shaping this zone. We investigated the population genetic structure of both species along a transect ranging from the core range of the expanding H. polyglotta across the contact zone and far into the range of the receding H. icterina. The theoretical predictions of changes in genetic diversity at the range edges were tested. No gradual change in genetic diversity was detected for both the expanding and the receding range margin. Furthermore, no genetic structure was found in either species supporting the hypothesis that long distance dispersal (LDD) occurs frequently due to the high mobility of these long‐distance migrants. The results suggest that when dispersal propensity is high and accompanied by frequent LDD events, then neither an enrichment nor a depletion of alleles along moving range edges would be detected. This these species as the probability to retain genetic diversity during exogenous induced range shifts is high in such mobile species.     

The spatio-temporal dynamics of neutral genetic diversity

The notions of pulled and pushed solutions of reaction-dispersal equations introduced by Garnier et al. (2012) and Roques et al. (2012) are based on a decomposition of the solutions into several components. In the framework of population dynamics, this decomposition is related to the spatio-temporal evolution of the genetic structure of a population. The pulled solutions describe a rapid erosion of neutral genetic diversity, while the pushed solutions are associated with a maintenance of diversity. This paper is a survey of the most recent applications of these notions to several standard models of population dynamics, including reaction-diffusion equations and systems and integro-differential equations. We describe several counterintuitive results, where unfavorable factors for the persistence and spreading of a population tend to promote diversity in this population. In particular, we show that the Allee effect, the existence of a competitor species, as well as the presence of climate constraints are factors which can promote diversity during a colonization. We also show that long distance dispersal events lead to a higher diversity, whereas the existence of a nonreproductive juvenile stage does not affect the neutral diversity in a range-expanding population.     

Testing Darwin's transoceanic dispersal hypothesis for the inland nettle family (Urticaceae)

Dispersal is a fundamental ecological process, yet demonstrating the occurrence and importance of long‐distance dispersal (LDD) remains difficult, having rarely been examined for widespread, non‐coastal plants. To address this issue, we integrated phylogenetic, molecular dating, biogeographical, ecological, seed biology and oceanographic data for the inland Urticaceae. We found that Urticaceae originated in Eurasia c. 69 Ma, followed by ≥ 92 LDD events between landmasses. Under experimental conditions, seeds of many Urticaceae floated for > 220 days, and remained viable after 10 months in seawater, long enough for most detected LDD events, according to oceanographic current modelling. Ecological traits analyses indicated that preferences for disturbed habitats might facilitate LDD. Nearly half of all LDD events involved dioecious taxa, so population establishment in dioecious Urticaceae requires multiple seeds, or occasional selfing. Our work shows that seawater LDD played an important role in shaping the geographical distributions of Urticaceae, providing empirical evidence for Darwin's transoceanic dispersal hypothesis.     

East-west divergence in central Brazilian Cerrado revealed by cpDNA sequences of a bird-dispersed tree species

Population genetic studies of plant species can contribute to understanding the evolutionary history of the Brazilian Cerrado, a highly threatened hotspot and the richest tropical savanna in the world. Therefore, this study aimed at investigating the diversity and genetic structure of Byrsonima coccolobifolia (Malpighiaceae), one of the most common tree species in the Brazilian Cerrado, focusing mainly on the central area of the biome. This is a bird-dispersed species, and studies targeting species with this seed dispersal syndrome are scarce. First, we performed a careful screening of cpDNA markers to identify regions suitable for intraspecific genetic studies, and then we investigated the diversity and genetic structure in ten populations of B. coccolobifolia. The cpDNA regions selected (trnS-trnG and trnH-trnK) revealed considerable divergence among populations and a striking geographical structure, separating populations in two groups, east and west. Although seed dispersal by birds is expected to reach long distances, our results strongly corroborate studies targeting trees with limited seed dispersal. This suggests that historical fragmentation of Cerrado (possibly during cold and dry periods of the Quaternary) might have limited long distance dispersal events after climate amelioration even in bird-dispersed species, especially while Cerrado was expanding but still fragmented.     

Long‐distance dispersal syndromes matter: diaspore–trait effect on shaping plant distribution across the Canary Islands

Oceanic islands emerge lifeless from the seafloor and are separated from continents by long stretches of sea. Consequently, all their species had to overcome this stringent dispersal filter, making these islands ideal systems to study the biogeographic implications of long‐distance dispersal (LDD). It has long been established that the capacity of plants to reach new islands is determined by specific traits of their diaspores, historically called dispersal syndromes. However, recent work has questioned to what extent such dispersal‐related traits effectively influence plant distribution between islands. Here we evaluated whether plants bearing dispersal syndromes related to LDD – i.e. anemochorous (structures that favour wind dispersal), thalassochorous (sea dispersal), endozoochorous (internal animal dispersal) and epizoochorous (external animal dispersal) syndromes – occupy a greater number of islands than those with unspecialized diaspores by virtue of their increased dispersal ability. We focused on the native flora of the lowland xeric communities of the Canary Islands (531 species) and on the archipelago distribution of the species. We controlled for several key factors likely to affect the role of LDD syndromes in inter‐island colonization, namely: island geodynamic history, colonization time and phylogenetic relationships among species. Our results clearly show that species bearing LDD syndromes have a wider distribution than species with unspecialized diaspores. In particular, species with endozoochorous, epizoochorous and thalassochorous diaspore traits have significantly wider distributions across the Canary archipelago than species with unspecialized and anemochorous diaspores. All these findings offer strong support for a greater importance of LDD syndromes on shaping inter‐island plant distribution in the Canary Islands than in some other archipelagos, such as Galápagos and Azores.     

Long-distance dispersal research: building a network of yellow brick roads

This special issue of Diversity and Distributions presents six papers that contribute to the assembly of a general research agenda for studying long‐distance dispersal (LDD) across a variety of taxonomic groups (e.g. birds, fish, aquatic invertebrates and plants), ecosystems (e.g. terrestrial and marine ecosystems, wetlands and grasslands) and thematic fields (e.g. biological transport, marine biology, biogeochemistry and biodiversity conservation). This editorial emphasizes the need to develop a network integrating different research approaches (‘yellow brick roads’) to address the great challenge (‘finding the end of the rainbow’) of quantifying, understanding and predicting LDD and its implications. I review the key avenues for future research suggested in the special issue contributions, and stress the critical importance of properly considering the spatial and temporal scales relevant to the process and system of interests. I propose combining absolute and proportional definitions of LDD as a default practice in any investigation of LDD processes. When LDD is defined primarily by an absolute critical distance that characterizes key feature(s) of the system of interest, a quantitative assessment of the proportion of dispersal events expected to move beyond this critical threshold distance should also be provided. When LDD is defined primarily by a certain small fraction of dispersal events that travel longer than all others, an estimate of the absolute distance associated with this high percentile at the tail of the dispersal curve should also be added.     

Pleistocene megafaunal extinctions and the functional loss of long‐distance seed‐dispersal services

Pleistocene extinctions affected mainly large‐bodied animals, determining the loss or changes in numerous ecological functions. Evidence points to a central role of many extinct megafauna herbivores as seed dispersers. An important step in understanding the legacy of extinct mutualistic interactions is to evaluate the roles and effectiveness of megafauna herbivores in seed dispersal. Here we use morphological and ecophysiological allometries to estimate both quantitative and qualitative aspects of seed‐dispersal services likely provided by extinct megafauna. We developed a mechanistic model that encompasses four stages of seed dispersal – seed ingestion, gut retention, animal movement, and seed deposition. We estimate seed‐dispersal kernels through simulations to infer the role of Pleistocene megafauna in promoting long‐distance dispersal and examine how seed dispersal was affected by extinctions. Simulations suggest extinct large‐bodied frugivores would frequently disperse large seeds over a thousand meters, whereas smaller‐bodied frugivores are more likely to deposit the seeds over a few hundred meters. Moreover, events of long‐distance seed dispersal by the extinct megafauna would be up to ten times longer than long‐distance dispersal by smaller‐sized extant mammals. By estimating the combined distribution of seed dispersal distances considering all large‐bodied mammalian frugivores in specific South American Pleistocene assemblages we found that long‐distance dispersal contracted by at least two thirds after the megafauna died out. The disruption of long‐distance dispersal is expected to have consequences for recruitment, spatial and genetic structure of plant populations, population persistence and community composition. Promoting long‐distance seed dispersal was one among other salient features of extinct Pleistocene megafauna that reveal their influence on natural ecosystems. Modeling the consequences of megafaunal extinctions can offer quantitative predictions on the consequences of ongoing defaunation to plant populations and ecological communities.     

Retention time variability as a mechanism for animal mediated long-distance dispersal

Long-distance dispersal (LDD) events, although rare for most plant species, can strongly influence population and community dynamics. Animals function as a key biotic vector of seeds and thus, a mechanistic and quantitative understanding of how individual animal behaviors scale to dispersal patterns at different spatial scales is a question of critical importance from both basic and applied perspectives. Using a diffusion-theory based analytical approach for a wide range of animal movement and seed transportation patterns, we show that the scale (a measure of local dispersal) of the seed dispersal kernel increases with the organisms' rate of movement and mean seed retention time. We reveal that variations in seed retention time is a key determinant of various measures of LDD such as kurtosis (or shape) of the kernel, thinkness of tails and the absolute number of seeds falling beyond a threshold distance. Using empirical data sets of frugivores, we illustrate the importance of variability in retention times for predicting the key disperser species that influence LDD. Our study makes testable predictions linking animal movement behaviors and gut retention times to dispersal patterns and, more generally, highlights the potential importance of animal behavioral variability for the LDD of seeds.     

The relative contributions of seed and pollen dispersal to gene flow and genetic diversity in seedlings of a tropical palm

Seed and pollen dispersal shape patterns of gene flow and genetic diversity in plants. Pollen is generally thought to travel longer distances than seeds, but seeds determine the ultimate location of gametes. Resolving how interactions between these two dispersal processes shape microevolutionary processes is a long‐standing research priority. We unambiguously isolated the separate and combined contributions of these two dispersal processes in seedlings of the animal‐dispersed palm Oenocarpus bataua to address two questions. First, what is the spatial extent of pollen versus seed movement in a system characterized by long‐distance seed dispersal? Second, how does seed dispersal mediate seedling genetic diversity? Despite evidence of frequent long‐distance seed dispersal, we found that pollen moves much further than seeds. Nonetheless, seed dispersal ultimately mediates genetic diversity and fine‐scale spatial genetic structure. Compared to undispersed seedlings, seedlings dispersed by vertebrates were characterized by higher female gametic and diploid seedling diversity and weaker fine‐scale spatial genetic structure for female gametes, male gametes and diploid seedlings. Interestingly, the diversity of maternal seed sources at seed deposition sites (N _em) was associated with higher effective number of pollen sources (N _ep), higher effective number of parents (N _e) and weaker spatial genetic structure, whereas seed dispersal distance had little impact on these or other parameters we measured. These findings highlight the importance maternal seed source diversity (N _em) at frugivore seed deposition sites in driving emergent patterns of fine‐scale genetic diversity and structure.     

Long‐distance dispersal maximizes evolutionary potential during rapid geographic range expansion

Conventional wisdom predicts that sequential founder events will cause genetic diversity to erode in species with expanding geographic ranges, limiting evolutionary potential at the range margin. Here, we show that invasive European starlings (Sturnus vulgaris) in South Africa preserve genetic diversity during range expansion, possibly as a result of frequent long‐distance dispersal events. We further show that unfavourable environmental conditions trigger enhanced dispersal, as indicated by signatures of selection detected across the expanding range. This brings genetic variation to the expansion front, counterbalancing the cumulative effects of sequential founding events and optimizing standing genetic diversity and thus evolutionary potential at range margins during spread. Therefore, dispersal strategies should be highlighted as key determinants of the ecological and evolutionary performances of species in novel environments and in response to global environmental change.     

Inferring long-distance dispersal and topographic barriers during post-glacial colonization from the genetic structure of red maple (Acer rubrum L.) in New England

Aim  This study aims to assess the role of long-distance seed dispersal and topographic barriers in the post-glacial colonization of red maple ( Acer rubrum L.) using chloroplast DNA (cpDNA) variation, and to understand whether this explains the relatively higher northern diversity found in eastern North American tree species compared with that in Europe.
Location  North-eastern United States.
Methods  The distribution of intraspecific cpDNA variation in temperate tree populations has been used to identify aspects of post-glacial population spread, including topographic barriers to population expansion and spread by long-distance seed dispersal. We sequenced c.  370 cpDNA base pairs from 221 individuals in 100 populations throughout the north-eastern United States, and analysed spatial patterns of diversity and differentiation.
Results  Red maple has high genetic diversity near its northern range limit, but this diversity is not partitioned by topographic barriers, suggesting that the northern Appalachian Mountains were not a barrier to the colonization of red maple. We also found no evidence of the patchy genetic structure that has been associated with spread by rare long-distance seed dispersal in previous studies.
Main conclusions  Constraints on post-glacial colonization in eastern North America seem to have been less stringent than those in northern Europe, where bottlenecks arising from long-distance colonization and topographic barriers appear to have strongly reduced genetic diversity. In eastern North America, high northern genetic diversity may have been maintained by a combination of frequent long-distance dispersal, minor topographic obstacles and diffuse northern refugia near the ice sheet.     

Genome‐wide SNP data reveal improved evidence for Antarctic glacial refugia and dispersal of terrestrial invertebrates

Antarctica is isolated, surrounded by the Southern Ocean and has experienced extreme environmental conditions for millions of years, including during recent Pleistocene glacial maxima. How Antarctic terrestrial species might have survived these glaciations has been a topic of intense interest, yet many questions remain unanswered, particularly for Antarctica's invertebrate fauna. We examine whether genetic data from a widespread group of terrestrial invertebrates, springtails (Collembola, Isotomidae) of the genus Cryptopygus, show evidence for long‐term survival in glacial refugia along the Antarctic Peninsula. We use genome‐wide SNP analyses (via genotyping‐by‐sequencing, GBS) and mitochondrial data to examine population diversity and differentiation across more than 20 sites spanning >950 km on the Peninsula, and from islands both close to the Peninsula and up to ~1,900 km away. Population structure analysis indicates the presence of strong local clusters of diversity, and we infer that patterns represent a complex interplay of isolation in local refugia coupled with occasional successful long‐distance dispersal events. We identified wind and degree days as significant environmental drivers of genetic diversity, with windier and warmer sites hosting higher diversity. Thus, we infer that refugial areas along the Antarctic Peninsula have allowed populations of indigenous springtails to survive in situ throughout glacial periods. Despite the difficulties of dispersal in cold, desiccating conditions, Cryptopygus springtails on the Peninsula appear to have achieved multiple long‐distance colonization events, most likely through wind‐related dispersal events.     

Comparing short and long‐distance dispersal: modelling and field case studies

Dispersal is a factor of great importance in determining a species spatial distribution. Short distance dispersal (SDD) and long distance dispersal (LDD) strategies yield very different spatial distributions. In this paper we compare spatial spread patterns from SDD and LDD simulations, contrast them with patterns from field data, and assess the significance of biological and population traits. Simulated SDD spread using an exponential function generates a single circular patch with a well‐defined invasion front showing a travelling‐wave structure. The invasive spread is relatively slow as it is restricted to reproductive individuals occupying the outer zone of the circular patch. As a consequence of this dispersal dynamics, spread is slower than spread generated by LDD. In contrast, the early and fast invasion of the entire habitat mediated by power law LDD not only involves a significantly greater invasion velocity, but also an entirely different habitat occupation. As newly dispersed individuals soon reach very distant portions of the habitat as well as the vicinity of the original dispersal focus, new growing patches are generated while the main patch increases its own growth absorbing the closest patches. As a consequence of both dispersal and lower density dependence, growth of the occupied area is much faster than with SDD. SDD and LDD also differ regarding pattern generation. With SDD, fractal patterns appear only in the border of the invasion front in SDD when competitive interaction with residents is included. In contrast, LDD patterns show fractality both in the spatial arrangements of patches as well as in patch borders. Moreover, values of border fractal dimension inform on the dispersal process in relation with habitat heterogeneity. The distribution of patch size is also scale‐free, showing two power laws characteristic of small and large patch sizes directly arising from the dispersal and reproductive dynamics. Ecological factors like habitat heterogeneity are relevant for dispersal, although its importance is greater for SDD, lowering the invasion velocity. Among the life history traits considered, adult mortality, the juvenile bank and mean dispersal distance are the most relevant for SDD. For LDD, habitat heterogeneity and changes in life history traits are not so relevant, causing minor changes in the values of the scale‐free parameters. Our work on short and long distance dispersal shows novel theoretical differences between SDD and LDD in invasive systems (mechanisms of pattern formation, fractal and scaling properties, relevance of different life history traits and habitat variables) that correspond closely with field examples and were not analyzed, at least in this degree of detail, by the previously existing models.