Phylogeography of cave pseudoscorpions in southern Australia

Aim  To investigate molecular phylogenetic divergence and historical biogeography of the cave-dwelling pseudoscorpion genus Protochelifer.
Location  Caves and nearby epigean habitats in southern Australia were sampled from western Victoria, Naracoorte Caves, Flinders Ranges, Kangaroo Island, Nullarbor Plain and south-west Western Australia.
Methods  Allozyme electrophoresis (57 individuals) and a 569-base-pair section of the mtDNA COI gene (22 individuals) were used to reconstruct phylogenetic relationships among four cave species and three epigean species from 13 locations.
Results  Phylogenetic reconstruction using the allozyme and mtDNA sequence data revealed a similar topology, showing recent speciation of several Protochelifer populations in caves from Naracoorte to the Nullarbor Plain. Naracoorte Caves contained a single species, Protochelifer naracoortensis , found in four separate caves, while all other cave species appear to be restricted to single caves.
Main conclusions  At a local scale, as indicated by the four Naracoorte caves, dispersal is thought to occur via micro- and mesocaverns, and possibly by phoresy using insect or bat vectors. With current data we are unable to determine if cavernicolous species of Protochelifer have arisen from a single cave colonization event followed by phoretic dispersal on bats to other caves, or multiple cave-invasion events from independent epigean ancestors. Genetic heterogeneity among Protochelifer populations from Nullarbor caves suggest that P. cavernarum , the only species presently recorded from this region, is likely to constitute a species complex requiring further study to fully resolve its relationships.     

Dispersal ability and host-plant characteristics influence spatial population structure of monophagous beetles

Abstract.  1. Dispersal plays an integral role in determining spatial population structure and, consequently, the long-term survival of many species. Theoretical studies indicate that dispersal increases with population density and decreasing habitat stability. In the case of monophagous insect herbivores, the stability of host-plant populations may influence their spatial population structure.
2. The tallgrass prairie in Iowa, U.S.A. is highly fragmented and most prairie insects face a landscape with fewer habitat patches and smaller host-plant populations than 150 years ago, potentially making dispersal between patches difficult. Some herbivores, however, use native plant species with weedy characteristics that have increased in abundance because of disturbances.
3. Mark–recapture data and presence–absence surveys were used to examine dispersal and spatial population structure of two monophagous beetles with host plants that exhibit different population stability and have responded differently to fragmentation of tallgrass prairie.
4. Chrysochus auratus Fabricius exhibits a patchy population structure and has relatively large dispersal distances and frequencies. Its host plant is variable locally in time and space, but is more abundant than 150 years ago. The other species, Anomoea laticlavia Forster, exhibits a metapopulation or non-equilibrium population structure and has relatively small dispersal distances and frequencies. Its host-plant populations are stable in time and space.
5. The results indicate that dispersal ability of monophagous beetles reflects the life-history dynamics of their host plants, but the spatial population structure exhibited today is strongly influenced by how the host plants have responded to the fragmentation process over both time and space.     

Dispersal traits linked to range size through range location, not dispersal ability, in Western Australian angiosperms

Aim  We examine the relative importance of seed dispersal mode in determining the range size and range placement in 524 species from six focal plant families (Agavaceae, Euphorbiaceae, Malvacaeae, Sapindaceae, Proteaceae and Fabaceae ( Acacia )).
Location  Western Australia.
Methods  Taxa were categorized by dispersal mode and life-form and their distributions modelled using maxent . Geographical range size was compared amongst dispersal mode, life-form and biome using phylogenetically independent contrasts. Geographical range placement was considered in a similar manner.
Results  Range size did not vary with dispersal mode (ant versus wind and vertebrate dispersal) or life-form, and instead varied primarily as a function of the biogeographical region in which a species was found. Range placement, however, did vary among dispersal modes, with the consequence that diversity of wind- and ant-dispersed plants increased with latitude while the diversity of vertebrate-dispersed plants was more evenly distributed.
Main conclusions  For the taxa studied, range sizes were a function of the biogeographical region in which species were found. Although differences in range size may exist among species differing in dispersal modes, they are likely to be far smaller than differences among species from different biogeographical regions. The trait most likely to affect species geographical range size, and hence rarity and risks associated with other threats, may simply be the geographical region in which that species has evolved.     

The community-wide and guild-specific effects of pubescence on the folivorous insects of manzanitas Arctostaphylos spp.

Abstract.  1. Insect communities on 26 species of manzanita Arctostaphylos spp. (Ericaceae) were sampled in order to examine the effects of variation in foliar pubescence traits on a community of folivorous insects. Manzanitas vary widely in pubescence density, length, and glandularity both within and between species.
2. Linear models were fitted and evaluated to determine whether pubescence traits are associated with the species richness and abundance of folivorous insects after accounting for the effects of other relevant habitat and host-plant related characteristics.
3. Pubescence traits were clearly associated with both community-wide and guild-specific variation in the structure of the folivorous insect community of manzanitas, however the effects of pubescence were manifested primarily as effects on the abundance of folivores not on species richness. The species richness of folivorous insects on manzanitas was not associated with pubescence density or length but was associated positively with glandularity.
4. The abundance of all guilds except leaf-mining insects was lower on manzanitas having longer pubescence. In contrast, the abundance of external-chewing insects was higher on plants having denser pubescence and on plants having glandular pubescence.
5. Overall, the results suggest that both longer pubescence and the amount of contact between an insect and pubescence act quantitatively to decrease the abundance of external-feeding guilds of folivorous insects. The abundance of species in internal-feeding guilds that oviposit directly on leaves is unrelated to foliar pubescence traits in the host plant.     

Searching for mechanisms of synchrony in spatially structured gamebird populations

1.  Time series data on five species of gamebird from the Dolomitic Alps were used to examine the relative importance of dispersal and common stochastic events in causing synchrony between spatially structured populations.
2.  Cross-correlation analysis of detrended time series was used to describe the spatial pattern of fluctuations in abundance, while standardized time series were used to describe both fluctuations and the trend in abundance. There were large variations in synchrony both within and between species and only weak negative relationships with distance.
3.  Species in neighbouring habitats were more likely to be in synchrony than species separated by several habitats. Species with similar density-dependent structure were more likely to be in synchrony.
4.  In order to estimate the relative importance of dispersal and environmental stochasticity, we modelled the spatial dynamics of each species using two different approaches. First, we used estimating functions and bootstrapping of time series data to calculate the relative importance of dispersal and stochastic effects for each species. Second, we estimated the intensity of environmental stochasticity from climatic records during the breeding season and then modelled the dispersal rate and dispersal distance for each species. The two models exhibited similar results for rock ptarmigan, black grouse, hazel grouse and rock partridge, while contrasting patterns were observed for capercaillie.
5.  The results suggest that environmental stochasticity plays the dominant role in synchronizing the fluctuations of these galliform species, although there will also be some dispersal between populations.     

The release of genetically modified grasses. Part 2: the influence of wind direction on pollen dispersal

 In part 1 an experiment was described for determining the extent of pollen dispersal from a Lolium source. The results were used to test Bateman’s pollen dispersal equations, which were found to be not particularly useful for describing variation in pollen deposition with distance. An improvement is suggested here which takes the influence of wind direction into account. For 11 of the 12 datasets the new equations fit significantly better than the original ones. Mean wind directions were used to produce 15 data subsets for testing Bateman’s equations for dispersal downwind of a pollen source. These equations fitted only 4 of the data subsets, all of which were collected from traps facing towards the pollen source. The usefulness of the model equations in estimating the importance of turbulence is brought into question. It is shown that models incorporating only distance and wind direction do not accurately describe pollen deposition. The amount of pollen deposited does not always decrease smoothly with increasing distance from the source. The variation in the amount of pollen deposited is probably influenced by several factors, including wind speed and turbulence. Received: 5 June 1996 / Accepted: 11 October 1996     

Genetic diversity, host-specificity and unusual phylogeography of a cryptic, host-associated species complex of gall-inducing scale insects

Abstract.  1. Australia has a unique and speciose gall-inducing scale insect fauna that is primarily associated with Myrtaceae. Much of the diversity is currently undescribed or uncharacterised.
2. This study concerns Apiomorpha munita (Hemiptera), a scale insect that induces characteristic four-horned galls on eucalypts of subgenus Symphyomyrtus and exhibits extraordinary karyotypic diversity (2n = 6 – 2n > 100). The three described subspecies of A. munita are each confined to hosts in different sections of Eucalyptus . Previous chromosomal data, however, cast doubt on the validity of the groupings, as two of the subspecies share multiple, different karyotypes (2n = 6, 20, 22, and 24).
3. Allozyme data were used to examine species delimitation, chromosome evolution, host associations and population structure in A. munita .
4. A cryptic-species radiation was revealed, with at least five taxa each restricted to a discrete set of host eucalypt species. This is consistent with host-associated speciation.
5. Karyotypic variation within A. munita partially fits the five distinct genetic groups, but there are additional chromosomal changes that are not accompanied by detected genetic differentiation.
6. The population structure of taxa within the A. munita species complex suggests that there are high levels of inbreeding, as would be expected for scale insects in which adult females are sessile. Some genotypes, however, are found over great distances (up to 1100 km). This is an unusual population structure because it combines low mobility and local differentiation with occasional long-distance dispersal, probably mediated by wind-dispersal of first-instar nymphs.     

Estimates of Gene Flow in Drosophila Pseudoobscura Determined from Nucleotide Sequence Analysis of the Alcohol Dehydrogenase Region

The genetic structure of Drosophila pseudoobscura populations was inferred from a nucleotide sequence analysis of a 3.4-kb segment of the alcohol dehydrogenase (Adh) region. A total of 99 isochromosomal strains collected from 13 populations in North and South America were used to determine if any population departed from a neutral model and to estimate levels of gene flow between populations. This study also included the nucleotide sequences from two sibling species, D. persimilis and D. miranda. We estimated the neutral mutation parameter, 4N mu, in synonymous and noncoding sites for 17 subregions of Adh in each of nine populations with sample sizes greater than three. The nucleotide diversity data in the nine populations was tested for departures from an equilibrium neutral model with two statistical tests. The Tajima and the Hudson, Kreitman, Aguade tests showed that each population fails to reject a neutral model. Tests for genetic differentiation between populations fail to show any population substructure among the North American populations of D. pseudoobscura. The nucleotide diversity data is consistent with direct and indirect measures of gene flow that show extensive dispersal between populations of D. pseudoobscura.     

Aphid dispersal flight disseminates fungal pathogens and parasitoids as natural control agents of aphids

Abstract.  1. Dispersal flight, a well-known strategy for aphids to locate suitable plants, was studied for its possible role in disseminating fungal pathogens and parasitoids as natural control agents of aphids by air captures in Hangzhou, China during 2001–2005. Up to 3183 migratory alates of green peach aphid Myzus persicae were captured from air using a yellow-plus-plant trap on the top platform of a six-storey building in an urbanised area, and individually reared in a laboratory for ≥ 7 days.
2. Among the captured alates, 28.9% survived on average for 2.5 days and then died from mycoses attributed to 10 species of obligate or non-obligate aphid pathogens. These were predominated by Pandora neoaphidis , which was causative of 80% of the mycosed alates. Another 4.4% survived for an average of 3.7 days, followed by mummification of Aphidius gifuensis (52.9%) and Diaeretella rapae (47.1%).
3. Numerous progeny colonies individually initiated by infected, parasitised, or healthy alates were monitored daily for 12 days, and fitted well to a logistic equation depicting the potential of their post-flight colonisation and fecundity. Both infected and parasitised alates from air were highly capable of initiating progeny colonies independently, although their potential fecundity was greatly reduced compared with that of healthy counterparts.
4. Our results confirmed that both obligate and non-obligate pathogens can be widespread with aphid dispersal flight, and demonstrated that parasitoids also took advantage of the host flight for their dispersal. This study provides new insights into aphid dispersal biology.     

Predictability of stream insect distributions is dependent on niche position,but not on biological traits or taxonomic relatedness of species

Species distributions can be analysed under two perspectives: the niche‐based approach, which focuses on species–environment relationships; and the dispersal‐based approach, which focuses on metapopulation dynamics. The degree to which each of these two components affect species distributions may depend on habitat fragmentation, species traits and phylogenetic constraints. We analysed the distributions of 36 stream insect species across 60 stream sites in three drainage basins at high latitudes in Finland. We used binomial generalised linear models (GLMs) in which the predictor variables were environmental factors (E models), within‐basin spatial variables as defined by Moran's eigenvector maps (M models), among‐basin variability (B models), or a combination of the three (E + M + B models) sets of variables. Based on a comparative analysis, model performance was evaluated across all the species using Gaussian GLMs whereby the deviance accounted for by binomial GLMs was fitted on selected explanatory variables: niche position, niche breadth, site occupancy, biological traits and taxonomic relatedness. For each type of model, a reduced Gaussian GLM was eventually obtained after variable selection (Bayesian information criterion). We found that niche position was the only variable selected in all reduced models, implying that marginal species were better predicted than non‐marginal species. The influence of niche position was strongest in models based on environmental variables (E models) or a combination of all types of variables (E + M + B models), and weakest in spatial autocorrelation models (M models). This suggests that species–environment relationships prevail over dispersal processes in determining stream insect distributions at a regional scale. Our findings have clear implications for biodiversity conservation strategies, and they also emphasise the benefits of considering both the niche‐based and dispersal‐based approaches in species distribution modelling studies.     

The structure of the aquatic insect community associated with intertidal pools on a New Jersey salt marsh

Abstract. 1. Landlocked intertidal pools (potholes) are one of the common aquatic habitats associated with eastern North American salt marshes.
2. Twenty species of insects, of which only eight were common, occurred in the pothole habitat. The harshness of the physical environment incurred in the pothole habitat is largely responsible for this depauperate insect community.
3. Spatial variation in the diversity of the insect community is best explained by differential densities of predaceous fish.
4. Seasonal changes in insect community diversity are attributable to fluctuations in water temperature and accompanying levels of dissolved oxygen.     

Genetic Variation for Dispersal by DROSOPHILA PSEUDOOBSCURA and DROSOPHILA PERSIMILIS

Release-recapture experiments using Drosophila pseudoobscura and D. persimilis strains of different karyotypes were performed in a heterogeneous environment. The heterogeneity was due to both spatial variation and the species of yeast used to attract the released flies. No karyotypic-specific habitat preferences were detected. However, in all releases, different strains did behave differently with respect to one or both of the heterogeneous factors. These results indicate there is variation for dispersal behavior in these species that is most likely based on genotype-dependent habitat preferences.     

Correlations between observed dispersal capabilities and patterns of genetic differentiation in populations of four aquatic insect species from the Arizona White Mountains, U.S.A.

SUMMARY 1. Dispersal ability is an important ecological factor that can influence population structure. In an attempt to determine the extent that the pattern of genetic differentiation is correlated with dispersal ability in stream-dwelling aquatic insects, we used the amplified fragment length polymorphism (AFLP) technique to characterise genetic variation in four aquatic insect species: Gumaga griseola (Trichoptera: Sericostomatidae), Helicopsyche mexicana (Trichoptera: Helicopsychidae), Psephenus montanus (Coleoptera: Psephenidae) and Ambrysus thermarum (Hemiptera: Naucoridae). Individuals were sampled from several sites within two adjacent catchments in the Arizona White Mountains. In addition to the genetic analyses, a 20-week-long trapping study was used to determine the relative dispersal ability of adults of the four species examined.
2. We obtained hierarchical indicators of genetic differentiation for catchments, sites within catchments and sites across the region examined. Overall, average estimators of genetic differentiation ( F -statistics) were consistent with direct observations of organismal movement, although it was our direct observations on adult insect flight that permitted us to interpret our results correctly. This was because of the fact that a lack of genetic differentiation across watersheds can be interpreted in two ways.
3. In contrast to F -statistics, patterns of genetic isolation by distance for each species more clearly reflected dispersal ability, suggesting that such analytical approaches provide less ambiguous information about the importance of gene flow in the hierarchical partitioning of genetic variation in stream organisms.     

Spatial and temporal variation in dispersal pattern of an invasive pine

Understanding dispersal ability of an invasive species is crucial for describing its potential spread. Despite this, we still know little about the dispersal potential of many invasive species. We explored dispersal spectra in Pinus strobus, an invasive tree in sandstone areas in Central Europe. We studied dispersal of the species using distribution of self-sown trees in the field. We compared these observed data with theoretical dispersal curves derived using information on wind speed, seed terminal velocity and tree height. Finally, we fitted various empirical dispersal curves to the observed data. All the analyses based on the observed field data were done for the whole dataset, and for the dataset divided by habitat types and age categories of the self-sown trees. P. strobus seeds can disperse up to 757.5 m from the source. The observed data fall within the confidence intervals of the predictions based on a negative exponential model. When comparing different dispersal functions fitted to the data, it was not easy to decide which of the dispersal curves provides the overall best fit. This was because different functions were the best predictors of different parts of the dispersal curve. We suggest that future studies should provide not only empirical fitted dispersal curves but also observed data and provide estimates of confidence intervals. Such information will provide insights into the reliability of the dispersal estimates in general and help to evaluate the predictive power of the different models.     

Progressive restriction of host plant exploitation along a climatic gradient: the willow psyllid Cacopsylla groenlandica in Greenland

1. Host plant selection by the endemic willow psyllid Cacopsylla groenlandica was studied at eight sites in three locations along a N–S climatic gradient in west Greenland.
2. C. groenlandica oviposited and developed on four willow species: Salix glauca, S. arctophila, S. uva-ursi and S. herbacea. Development took place on female catkins of all species and on leaf shoots of S. glauca .
3. At sites in southern Greenland, C. groenlandica fully utilized all four willow species. However, there was progressive reduction of host plant species and tissues exploited as latitude increased, such that towards the northern limit of its range the psyllid became highly specialized, feeding only on female catkins of S. glauca , despite the presence of alternative hosts. Where plant growth was tightly constrained within a restricted growing season, the demands of synchronizing with more than one species may have become too great, resulting in specialization on a single host.
4. The probable mechanisms producing specialization were between-species differences in host plant phenology, linked to variation in the growing season, set against the partially non-overlapping geographical distribution patterns of the host willows.
5. A small degree of phenological asynchrony between host plants can be advantageous for the insect. Under favourable conditions the reproductive season can be extended by using a series of host plants with staggered phenologies. Some 'early' hosts also create predator-free space. However, as asynchrony increases, the advantages disappear as the insect becomes unable to exploit the full range of host options.
6. The importance of host plant synchrony in the life histories of arctic insects is discussed.     

Modelling pest population resurgence due to recolonization of fields following an insecticide application

1. To investigate the effect of insect recolonization on the insecticide-induced resurgence of crop pests, a modified Lotka–Volterra predator–prey model was partitioned into two areas, 'sprayed' and 'unsprayed'.
2. The unsprayed area provided a source of insects to recolonize the sprayed area, resulting in a change in pest dynamics in both areas following an insecticide application.
3. Model sensitivity to insecticide selectivity and rates of predation, insect dispersal and pest population increase were examined.
4. Resurgence risk in the sprayed area increased with increasing pest dispersal rate, but decreased with increasing predator dispersal rate.
5. Pest resurgence could also occur in the unsprayed area, especially when prey dispersal rates were low. The extent of resurgence in the unsprayed area could in some circumstances be worse than in the sprayed area itself.
6. The more efficient and longer-lived the predators, the greater the level of pest resurgence in both areas following insecticide use.
7. More selective insecticides, killing the pest, but not the predator, reduced resurgence provided that the average life-span of the predators was reasonably long. Even highly selective insecticides could cause resurgence of the pest in the unsprayed area.
8. When the prey carrying capacity of the unsprayed area was increased relative to that of the sprayed area, resurgent effects in the unsprayed area were reduced, but could still be significant under some circumstances.     

Genetic structure and dispersal of Macrobrachium australiense (Decapoda: Palaemonidae) in western Queensland, Australia

SUMMARY 1. The freshwater prawn, Macrobrachium australiense (Decapoda: Palaemonidae), is an abundant species throughout the rivers of western Queensland, and it is thought to have effective dispersal capabilities. Given the very low topographical relief of the western Queensland landscape and the vast nature of episodic flooding, it was predicted that widespread dispersal in this species would occur within, and possibly between the four major catchments of the region: the Darling, Bulloo, Cooper and Diamantina. We analysed eight polymorphic allozyme systems and a fragment of the cytochrome c oxidase subunit 1 (COI) mitochondrial DNA (mtDNA) gene to determine the extent of recent and historical patterns of dispersal at nested spatial scales, within and between catchments.
2. Large and significant levels of allozyme and mtDNA differentiation were revealed among all catchments, indicating that dispersal of M. australiense does not occur across catchment boundaries, although this species is reportedly capable of overland movement. In contrast, no significant patterns of genetic differentiation were resolved between major subcatchments of the Darling and Cooper, or between sites within these subcatchments, indicating that populations of M. australiense are panmictic within catchments.
3. The MtDNA data resolved two divergent and reciprocally monophyletic clades, with the first representing the Darling catchment, and the second corresponding to the Bulloo, Cooper and Diamantina catchments. We postulate that extreme variation in historical climatic patterns and palaeohydrologic conditions played an important role in shaping the population structure of M. australiense throughout western Queensland during the Quaternary.     

Arthropod faunal similarity of Bornean rain forest trees

ABSTRACT. 1. Arthropod samples, collected by insecticide fogging ten Bornean lowland rain forest trees, were sorted to species and used to study some of the determinants of faunal similarity of these trees.
2. Similarity indices were calculated between pairs of trees for guilds of beetles, beetle families and for different orders and other families of insects.
3. Multiple regression analyses of five independent variables of the trees (taxonomic similarity, distance apart, vertical canopy overlap, similarity in tree height, epiphytic load) were carried out against these similarity indices.
4. In eight of the eighteen insect groups analysed, taxonomic similarity of the trees is the most important variable affecting their species composition, accounting for a maximum of 29.4% of the variation.
5. For the Homoptera, Gryllidae, Anthicidae, Chrysomeloidea and scavengers, distance between the trees and the similarity of the trees'epiphyte load have a greater effect on their faunal similarities than taxonomic similarity of the trees.
6. The maximum variation accounted for by variables with significant T ratios was 38.6% in the Anthicidae. In four groups, the Formicidae, Heteroptera, Galerucinae and Corylophidae, none of the variables was significant.
7. These data may support the view that for many insect groups there is less host-specificity in the tropics than in temperate regions.     

Evaluation of the genomic extent of effects of fixed inversion differences on intraspecific variation and interspecific gene flow in Drosophila pseudoobscura and D. persimilis

There is increasing evidence that chromosomal inversions may facilitate the formation or persistence of new species by allowing genetic factors conferring species-specific adaptations or reproductive isolation to be inherited together and by reducing or eliminating introgression. However, the genomic domain of influence of the inverted regions on introgression has not been carefully studied. Here, we present a detailed study on the consequences that distance from inversion breakpoints has had on the inferred level of gene flow and divergence between Drosophila pseudoobscura and D. persimilis. We identified the locations of the inversion breakpoints distinguishing D. pseudoobscura and D. persimilis in chromosomes 2, XR, and XL. Population genetic data were collected at specific distances from the inversion breakpoints of the second chromosome and at two loci inside the XR and XL inverted regions. For loci outside the inverted regions, we found that distance from the nearest inversion breakpoint had a significant effect on several measures of divergence and gene flow between D. pseudoobscura and D. persimilis. The data fitted a logarithmic relationship, showing that the suppression of crossovers in inversion heterozygotes also extends to loci located outside the inversion but close to it (within 1-2 Mb). Further, we detected a significant reduction in nucleotide variation inside the inverted second chromosome region of D. persimilis and near one breakpoint, consistent with a scenario in which this inversion arose and was fixed in this species by natural selection.     

Detecting dispersal of Nuphar lutea in river corridors using microsatellite markers

1. River corridors are well-known for their role in plant dispersal. The buoyancy of seeds, the possibility of dispersal by vegetative fragments, and the frequency and efficiency of dispersal among different river catchments determine linear distribution patterns. Little is known about the relative importance of these factors to observed patterns of genetic variation.
2. One hundred and fifty-six Nuphar lutea individuals from forty-four sampling sites in the river catchments of the Cidlina River, the Mrlina River and the Labe River (Czech Republic) were studied using ten microsatellite markers. Interpretation of patterns in genetic variation allowed several conclusions about dispersal mechanisms.
3. Vegetative long-distance dispersal is probably very limited in this species. Only one multilocus genotype was found in more than one sampling site. The distance between the sites was about 75 km.
4. To explain the distribution of Bayesian based clusters of related multilocus genotypes, both along-river and inter-river long-distance dispersals have to be invoked.
5. A marginally significant tendency for higher genetic diversity in the lower part of the river Cidlina was detected. Continuous downstream dispersal of seeds by water currents could be a valid explanation.
6. Significant positive autocorrelation was found among individuals at within-river distances of up to 25 km. Repeated dispersal of seeds over distances in the range of tens of kilometers is common.