Increases in air temperature can promote wind-driven dispersal and spread of plants

Long-distance dispersal (LDD) of seeds and pollen shapes the spatial dynamics of plant genotypes, populations and communities. Quantifying LDD is thus important for predicting the future dynamics of plants exposed to environmental changes. However, environmental changes can also alter the behaviour of LDD vectors: for instance, increasing air temperature may enhance atmospheric instability, thereby altering the turbulent airflow that transports seed and pollen. Here, we investigate temperature effects on wind dispersal in a boreal forest using a 10-year time series of micrometeorological measurements and a Lagrangian stochastic model for particle transport. For a wide range of dispersal and life history types, we found positive relations between air temperature and LDD. This translates into a largely consistent positive effect of +3°C warming on predicted LDD frequencies and spread rates of plants. Relative increases in LDD frequency tend to be higher for heavy-seeded plants, whereas absolute increases in LDD and spread rates are higher for light-seeded plants for which wind is often an important dispersal vector. While these predicted increases are not sufficient to compensate forecasted range losses and environmental changes can alter plant spread in various ways, our results generally suggest that warming can promote wind-driven movements of plant genotypes and populations in boreal forests.     

种子的长距离风传播模型研究进展

 植物种子的长距离传播在物种迁移、生物入侵、保护生物学等领域有重要的生态和进化意义。种子传播有很多方式,开阔草原等地区的草本植物和许多热带和温带的树木都是通过风传播种子的。风传播的方式最适合进行种子长距离传播现象的模拟研究。种子的风传播模型是传播生态研究的一个重要领域,尤其是种子的长距离风传播模型,对于外来入侵植物的扩散和破碎化景观中植物种群的基因交流等生态过程研究举足轻重,然而国内鲜见这方面的研究成果。本文综述了种子长距离风传播现象研究的背景和意义,分析了风传播种子模型的基本形式和构成原理,并分别就现象模型和机理模型的相关研究进展进行了总结,同时指出了未来发展的几个重要方向。种子的风传播模型可以分为现象模型和机理模型两类,现象模型按种子传播核心的形式包括短尾模型、偏峰长尾模型和混合传播核心模型,后两者对于长距离传播数据的模拟可以取得很好的效果。机理模型按照模拟机制可分为欧拉对流扩散模型和拉格郎日随机模型两类。本文重点介绍了种子的长距离风传播现象的形成机理和两类机理模型的参数构成和处理方式。适合种子脱落的天气和适合传播的天气的同步性可能是形成种子长距离风传播的一个重要前提,林缘和地表存在的上升气流及大风和暴风中形成的速度梯度都可能对于种子的长距离传播有重要的作用。机理模型的操作因子主要包括生物方面的因子、气象方面的因子和地形方面的因子。同时对目前几个应用比较成功的机理模型进行了简要的介绍和评价,包括倾斜羽毛模型、对流-扩散-下降模型、无掩蔽模型、背景模型、WINDISPER及其改进模型和PAPPUS模型。最后指出,目前在风传播种子的长距离模型研究中,对草本植物种子的传播模拟的投入明显不如树木种子的长距离传播模拟,对于破碎化景观中种子长距离的风传播的研究还存在很大的差距,而对提高机理模型预测能力的高分辨率物理环境数据输入技术的需求则为多学科交叉提供了很好的机会。     

风传草本植物种子空间传播新模型

种子阶段产生的时空格局对植物种群迁移十分关键,在植物种群动态、外来种入侵、保护生物学领域有重要意义。大多数种子从母株处传播距离较短,少数种子可以进行长距离传播。近30多年对种子传播的大量研究集中在近距离传播,这与目前的技术很难直接跟踪长距离传播的种子有关,尤其是草本植物种子传播数据极难获取。在现实条件约束下,利用模型模拟种子传播显示了重要作用,但目前尚未形成种子密度随距离变化的统一性函数或分布曲线,我国更是鲜见此方面研究成果。提出了一种全新的风传草本植物种子的空间传播模型,即准麦克斯韦空间分布模型,将种子长、中、短距离的传播归纳于统一的模型之中;实现无风或有风时,任意方向及区域的种子传播模拟,并且利用前人已有的实验或实测数据对模型进行了适应性分析,旨在推动此领域研究的进一步探索与发展。     

Should I stay or should I go? Mycorrhizal plants are more likely to invest in long‐distance seed dispersal than non‐mycorrhizal plants

Seed dispersal and mycorrhizal associations are key mutualisms for the functioning and regeneration of plant communities; however, these processes have seldom been explored together. We hypothesised that obligatory mycorrhizal plants will be less likely to have long‐distance dispersal (LDD) syndromes since the probability of finding suitable mycorrhizal partners is likely to decrease with distance to the mother plant. We contrasted the mycorrhizal status and LDD syndromes for 1960 European plant species, using phylogenetically corrected log‐linear models. Contrary to our expectation, having specialised structures for LDD is more frequent in obligate mycorrhizal plants than in non‐mycorrhizal plants, revealing that lack of compatible mutualists does not constrain investment in LDD structures in the European Flora. Ectomycorrhizal plants associated with wind‐dispersing fungi are also more likely to have specialised structures for wind dispersal. Habitat specificity and narrower niche of non‐mycorrhizal plants might explain the smaller investment in specialised structures for seed dispersal.     

Dandelion Seed Dispersal: The Horizontal Wind Speed Does Not Matter for Long-Distance Dispersal - it is Updraft!

Abstract: Long-distance dispersal of seeds (LDD) surely affects most ecological and evolutionary processes related to plant species. Hence, numerous attempts to quantify LDD have been made and, especially for wind dispersal, several simulation models have been developed. However, the mechanisms promoting LDD by wind still remain ambiguous and the effects of different weather conditions on LDD, although recognized as important, have only rarely been investigated. Here we examine the influence of wind speed and updrafts on dispersal of dandelion ( Taraxacum officinale agg.), a typical wind-dispersed herb of open habitats. We used PAPPUS, a weather-sensitive mechanistic simulation model of wind dispersal, which considers frequency distribution of weather conditions during the period the simulation refers to. A simulation for the 4-month shedding period of dandelion shows that high wind speed does not promote LDD. In contrast, vertical turbulence, especially convective updrafts, are of overwhelming importance. Mainly caused by updrafts, in the simulations more than 0.05 % of dandelion seeds were dispersed beyond 100 m, a distance commonly used to define LDD. We conclude that long-distance dispersal of seeds of herbaceous species with falling velocities < 0.5 - 1.0 ms^-1 is mainly caused by convective updrafts.     

Overseas seed dispersal by migratory birds

Long-distance dispersal (LDD) promotes the colonization of isolated and remote habitats, and thus it has been proposed as a mechanism for explaining the distributions of many species. Birds are key LDD vectors for many sessile organisms such as plants, yet LDD beyond local and regional scales has never been directly observed nor quantified. By sampling birds caught while in migratory flight by GPS-tracked wild falcons, we show that migratory birds transport seeds over hundreds of kilometres and mediate dispersal from mainland to oceanic islands. Up to 1.2% of birds that reached a small island of the Canary Archipelago (Alegranza) during their migration from Europe to Sub-Saharan Africa carried seeds in their guts. The billions of birds making seasonal migrations each year may then transport millions of seeds. None of the plant species transported by the birds occurs in Alegranza and most do not occur on nearby Canary Islands, providing a direct example of the importance of environmental filters in hampering successful colonization by immigrant species. The constant propagule pressure generated by these LDD events might, nevertheless, explain the colonization of some islands. Hence, migratory birds can mediate rapid range expansion or shifts of many plant taxa and determine their distribution.     

Migratory strategies of waterbirds shape the continental‐scale dispersal of aquatic organisms

Long distance dispersal (LDD) of propagules is an important determinant of population dynamics, community structuring and biodiversity distribution at landscape, and sometimes continental, scale. Although migratory animals are potential LDD vectors, migratory movement data have never been integrated in estimates of propagule dispersal distances and LDD probability. Here we integrated migratory movement data of two waterbird species (mallard and teal) over two continents (Europe and North America) and gut retention time of different propagules to build a simple mechanistic model of passive dispersal of aquatic plants and zooplankton. Distance and frequency of migratory movements differed both between waterbird species and continents, which in turn resulted in changes in the shapes of propagule dispersal curves. Dispersal distances and the frequency of LDD events (generated by migratory movements) were mainly determined by the disperser species and, to a lesser extent, by the continent. The gut retention time of propagules also exerted a significant effect, which was mediated by the propagule characteristics (e.g. seeds were dispersed farther than Artemia cysts). All estimated dispersal curves were skewed towards local‐scale dispersal and, although dispersal distances were lower than previous estimates based only on the vector flight speed, had fat tails produced by LDD events that ranged from 230 to 1209 km. Our results suggest that propagule dispersal curves are determined by the migratory strategy of the disperser species, the region (or flyway) through which the disperser population moves, and the propagule characteristics. Waterbirds in particular may frequently link wetlands separated by hundreds of kilometres, contributing to the maintenance of biodiversity and, given the large geographic scale of the dispersal events, to the readjustment of species distributions in the face of climate change.     

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.     

The establishment of plants following long-distance dispersal

Long-distance dispersal (LDD) beyond the range of a species is an important driver of ecological and evolutionary patterns, but insufficient attention has been given to postdispersal establishment. In this review, we summarize current knowledge of the post-LDD establishment phase in plant colonization, identify six key determinants of establishment success, develop a general quantitative framework for post-LDD establishment, and address the major challenges and opportunities in future research. These include improving detection and understanding of LDD using novel approaches, investigating mechanisms determining post-LDD establishment success using mechanistic modeling and inference, and comparison of establishment between past and present. By addressing current knowledge gaps, we aim to further our understanding of how LDD affects plant distributions, and the long-term consequences of LDD events.     

The role of migratory ducks in the long‐distance dispersal of native plants and the spread of exotic plants in Europe

Little is known about the role of migratory waterfowl in the long‐distance dispersal (LDD) of seeds. We studied the gut contents of 42 teals Anas crecca collected in the Camargue, southern France, and found intact seeds of 16 species. There was no relationship between the probability that a given seed species was found intact in the lower gut, and the seed hardness or size. The number of seeds found in the oesophagus and gizzard (a measure of ingestion rate) was the only significant predictor of the occurrence of intact seeds in the lower gut, so studies of waterfowl diet can be used as surrogates of dispersal potential. In a literature review, we identified 223 seed species recorded in 25 diet studies of teal, pintail Anas acuta, wigeon A. penelope or mallard A. platyrhynchos in Europe. We considered whether limited species distribution reduces the chances that a seed can be carried to suitable habitat following LDD. Overall, 72% of plant species recorded in duck diets in southern Europe (36 of 50) were also recorded in the north, whereas 97% of species recorded in duck diets in the north (137 of 141) were also recorded in the south. This suggests a great potential for LDD, since most dispersed plants species occur throughout the migratory range of ducks. Migratory ducks are important vectors for both terrestrial and aquatic plant species, even those lacking the fleshy fruits or hooks typically used to identify seeds dispersed by birds. Finally, we show ducks are important vectors of exotic plant species. We identified 14 alien to Europe and 44 native to Europe but introduced to some European countries whose seeds have been recorded in duck diet.     

Importance of functional traits and regional species pool in predicting long-distance dispersal in savanna ecosystems

Long-distance dispersal (LDD) of plants is difficult to measure but disproportionately important for various ecological and evolutionary processes. Dispersal of seeds of gallery-forest trees in savanna provides an opportunity for the study of colonisation processes and species coexistence driven by LDD. Investigations were carried out on 91 isolated trees along four gallery forests sampled in the Biosphere Reserve of Pendjari, Benin. The abundance of adult trees within nearest gallery forest was combined with functional traits (species maximum height, seed weight, morphological adaptation for dispersal by wind, water, birds and mammals) to explain the floristic composition of forest seedlings and saplings under isolated trees and in savanna. Stepwise negative binomial regression was used to identify the most significant variables explaining abundance of seedlings and saplings beneath isolated trees and in savanna and then derive colonisation from seedlings and persistence from saplings. The maximum height of species and seed weight explained the highest proportion of variance in species colonisation. Morphological dispersal syndromes by wind and birds had poor explanatory importance. Species rare in gallery forest had higher potential to colonise new environments through LDD whilst abundant species had higher persistence abilities. Contrary to the predictions of the seedling-size effect, small-seeded species dominated the sapling stage. The findings revealed the strong dependence of LDD and subsequent colonisation and persistence processes on species traits specialised for a variety of dispersal vectors. They also suggest that LDD towards isolated trees established far away from gallery forest can be difficult.     

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.     

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.     

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.     

Recent advances in the study of long-distance dispersal of aquatic invertebrates via birds

Although Darwin pioneered the study of long‐distance dispersal (LDD) of aquatic invertebrates via waterbirds, it remains in its infancy as a modern discipline. A handful of recent studies have quantified internal or external transport in the field, confirming that a variety of long‐distance migrants carry invertebrates both internally and externally. These studies show that variation in the morphology of vectors influences the frequency and size of propagules transported, and suggest that more invertebrate groups disperse via birds than was previously thought. Dispersal limitation has mainly been investigated for zooplankton in small experimental systems from which waterbirds were effectively excluded, and the extent of such limitation for invertebrate populations in wetlands interconnected by waterbird movements remains unclear. We expect that the spatial and temporal scales at which dispersal limitation constrains geographical ranges, species richness and genetic structure of invertebrates depends partly on the density of migratory birds using the area. Birds may have a major role in the expansion of exotic species. We propose several avenues for future research. There is a particular need for more quantitative studies of LDD by birds that will enable modellers to assess its role in maintaining invertebrate biodiversity among increasingly fragmented wetlands and in the face of climate change, as well as in the spread of invasive species.     

Long-distance seed dispersal by wind: disentangling the effects of species traits,vegetation types,vertical turbulence and wind speed

Long-distance dispersal (LDD) of plant seeds by wind is affected by functional traits of the species, specifically seed terminal velocity and height of seed release above the vegetation cover (HAC), as well as by the meteorological parameters wind speed and vertical turbulence. The relative importance of these parameters is still under debate and the importance of their variability in vegetation types, sites and years has only rarely been quantified. To address these topics, we performed simulation studies for different vegetation types, sites, years and plant species with PAPPUS, a process based trajectory model. We found that LDD (measured in terms of migration rates) was higher in forests compared to open landscapes. Forests also showed greater between-year variability in LDD. Terminal velocity had an effect on LDD in both vegetation types, while the effect of HAC was significant only in the open landscape. We found considerable differences in how vertical turbulence and wind speed affect LDD between species and vegetation types: In the open landscape the strength of the positive relationship between vertical turbulence and LDD generally decreases with terminal velocity, whereas it increases in forests. The strength of the predominantly positive effect of wind speed on LDD increases with terminal velocity in both vegetation types, while in forests we found even negative relationships for species with low terminal velocity. Our results generally suggest that the effects of vertical turbulence and wind speed on LDD by wind diverge for species with different functional traits as well as in different vegetation types.     

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.     

Worldwide long‐distance dispersal favored by epizoochorous traits in the biogeographic history of Omphalodeae (Boraginaceae)

Biogeographic dispersal is supported by numerous phylogenetic results. In particular, transoceanic dispersal, rather than vicariance, is suggested for some plant lineages despite current long distances between America and Europe. However, few studies on the biogeographic history of plants have also studied the role of diaspore syndromes in long‐distance dispersal (LDD). Species of the tribe Omphalodeae (Boraginaceae) offer a suitable study system because the species have a wide variety of diaspore traits related to LDD and different lineages conform to patched worldwide distributions on three distant continents (Europe, America and New Zealand). Our aim is to reconstruct the biogeographical history of the Omphalodeae and to investigate the role of diaspore traits favoring LDD and current geographic distributions. To this end, a time‐calibrated phylogeny with 29 of 32 species described for Omphalodeae was reconstructed using biogeographical analyses (BioGeoBEARS, Lagrange) and models (DEC and DIVA) under different scenarios of land connectivity. Character‐state reconstruction (SIMMAP) and diversification rate estimations of the main lineages were also performed. The main result is that epizoochorous traits have been the ancestral state of LDD syndromes in most clades. An early diversification age of the tribe is inferred in the Western Mediterranean during late Oligocene. Colonization of the New World by Omphalodeae, followed by fast lineage differentiation, took place sometime in the Oligocene‐Miocene boundary, as already inferred for other angiosperm genera. In contrast, colonization of remote islands (New Zealand, Juan Fernández) occurred considerably later in the Miocene‐Pliocene boundary.     

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.     

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.