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.     

Characterization of 24 microsatellite markers in 11 species of fire ants in the genus Solenopsis (Hymenoptera: Formicidae)

The social parasite ant Solenopsis daguerrei infests colonies of several mound-building fire ant species. Twenty-four microsatellite markers were isolated from a repeat-enriched genomic library of S. daguerrei. Eleven loci were polymorphic in this ant with two to six alleles per locus. Expected heterozygosity ranged from 0.0222 to 0.7940 among loci. Most microsatellites amplified successfully across the 11 Solenopsis species tested and will be useful for evolutionary genetic studies in this diverse ant group.     

Propagule pressure and colony social organization are associated with the successful invasion and rapid range expansion of fire ants in China

We characterized patterns of genetic variation in populations of the fire ant Solenopsis invicta in China using mitochondrial DNA sequences and nuclear microsatellite loci to test predictions as to how propagule pressure and subsequent dispersal following establishment jointly shape the invasion success of this ant in this recently invaded area. Fire ants in Wuchuan (Guangdong Province) are genetically differentiated from those found in other large infested areas of China. The immediate source of ants in Wuchuan appears to be somewhere near Texas, which ranks first among the southern USA infested states in the exportation of goods to China. Most colonies from spatially distant, outlying areas in China are genetically similar to one another and appear to share a common source (Wuchuan, Guangdong Province), suggesting that long‐distance jump dispersal has been a prevalent means of recent spread of fire ants in China. Furthermore, most colonies at outlier sites are of the polygyne social form (featuring multiple egg‐laying queens per nest), reinforcing the important role of this social form in the successful invasion of new areas and subsequent range expansion following invasion. Several analyses consistently revealed characteristic signatures of genetic bottlenecks for S. invicta populations in China. The results of this study highlight the invasive potential of this pest ant, suggest that the magnitude of international trade may serve as a predictor of propagule pressure and indicate that rates and patterns of subsequent range expansion are partly determined by the interplay between species traits and the trade and transportation networks.     

Seed longevity and fire: germination responses of an exotic perennial herb in NW Patagonian grasslands (Argentina)

Fire affects grassland composition by selectively influencing recruitment. Some exotic species can increase their abundance as a consequence of fire-stimulated seed germination, but response may depend on seed age. Rumex acetosella L. (Polygonaceae, sheep's sorrel) is a cosmopolitan herb that has invaded NW Patagonia's grasslands. This species forms persistent soil seed banks and increases after disturbances, particularly fire. We studied how fire and seed longevity influence R. acetosella germination. In 2008, we conducted laboratory experiments where we exposed different-aged seeds (up to 19 years old) to heat, smoke, charcoal, ash and control treatments. Total percentage germination and mean germination time depended on both seed age and fire treatment. Germination of younger seeds decreased with increasing temperature. There was no general pattern in germination responses of different-aged seeds to smoke, charcoal and ash. While smoke improved the germination of fresh seeds, charcoal decreased germination. Germination of untreated seeds was negatively correlated with seed age, and mean germination time increased with seed age. In most treatments, fresh seeds had lower germination than 1-5-year-old seeds, indicating an after-ripening requirement. Smoke stimulates R. acetosella germination, causing successful recruitment during post-fire conditions. Fresh seeds are particularly responsive to fire factors, possibly because they have not experienced physical degradation and are more receptive to environmental stimuli. Knowing the colonisation potential from the soil seed bank of this species during post-fire conditions will allow us to predict their impact on native communities.     

Molecular evolutionary analyses of the odorant-binding protein gene Gp-9 in fire ants and other Solenopsis species

The fire ant Solenopsis invicta exists in two social forms, one with colonies headed by a single reproductive queen (monogyne form) and the other with colonies containing multiple queens (polygyne form). This variation in social organization is associated with variation at the gene Gp-9, with monogyne colonies harboring only the B allelic variant and polygyne colonies containing b-like variants as well. We generated new Gp-9 sequences from 15 Solenopsis species and combined these with previously published sequences to conduct a comprehensive, phylogenetically based study of the molecular evolution of this important gene. The exon/intron structure and the respective lengths of the five exons of Gp-9 are identical across all species examined, and we detected no evidence for intragenic recombination. These data conform to a previous suggestion that Gp-9 lies in a genomic region with low recombination, and they indicate that evolution of the coding region in Solenopsis has involved point substitutions only. Our results confirm a link between the presence of b-like alleles and the expression of polygyny in all South American fire ant species known to possess colonies of both social forms. Moreover, phylogenetic analyses show that b-like alleles comprise a derived clade of Gp-9 sequences within the socially polymorphic species, lending further support to the hypothesis that monogyny preceded polygyny in this group of fire ants. Site-specific maximum likelihood tests identified several amino acids that have experienced positive selection, two of which are adjacent to the inferred binding-pocket residues in the GP-9 protein. Four other binding-pocket residues are variable among fire ant species, although selection is not implicated in this variation. Branch-specific tests revealed strong positive selection on the stem lineage of the b-like allele clade, as expected if selection drove the amino acid replacements crucial to the expression of polygyne social organization. Such selection may have operated via the ligand-binding properties of GP-9, as one of the two amino acids uniquely shared by all b-like alleles is predicted to be a binding-pocket residue.     

Dynamic expansion in recently introduced populations of fire ant parasitoids (Diptera: Phoridae)

Combating invasive species requires a detailed, mechanistic understanding of the manner and speed with which organisms expand their ranges. Biological control efforts provide an opportunity to study the process of species invasions and range expansions under known initial conditions. This study examines the rate, pattern and mechanisms of spread for two populations of the biological control agent Pseudacteon tricuspis, phorid-fly parasitoids of imported fire ants. We employ a trap-based survey method that detects phorid flies in low-density populations, and provides data on abundance. This technique allows us to differentiate between continuous population spread and effective long-distance dispersal and to examine density gradients of phorid flies across the expanding population front. We find that occupied sites in front of the leading edge of continuous populations were common; forming small populations we refer to as satellite populations. Satellite populations are tens of kilometers from the nearest possible source. Wind governs the dynamics of spread in these two central Texas populations. Population edges expanding with the wind exhibited a higher frequency of effective long-distance dispersal than did populations expanding into the wind. This enhanced effective long-distance dispersal rate translated into a five times faster rate of spread for population edges traveling with the wind. This planned invasion shares many characteristics in common with unplanned species invasions including: protracted establishment phase during which densities were below detection thresholds, and slow initial spread immediately after establishment followed by rapid, accelerating spread rates as population sizes grew.     

Seed longevity and fire: post‐germination responses of Rumex acetosella L. in northwest Patagonian grasslands (Argentina)

The cosmopolitan herb Rumex acetosella forms persistent soil seed banks and increases in cover after fire. We investigated how the interaction between seed age and fire affects seedling growth by exposing different‐aged seeds to heat, smoke, charcoal, and ash treatments. We measured growth of germinated seedlings that were transplanted and allowed to grow for 65 days in a greenhouse. Seedlings from seeds >8 years old did not reach an appropriate radicle length for transplantation. Seedling growth decreased with increasing temperature of the heat treatment. As seed age increased, growth decreased with smoke and charcoal, and increased with ash treatment. Height was negatively correlated with seed age. Our results suggest that fire and seed age could affect demographic responses of R. acetosella seedling populations. Post‐fire recruitment could be partially favored by the positive effect of nutrient input from ash on seedling growth. High fire intensities, however, would be detrimental to seedling vigor.     

Population dynamics of the invasive Pacific oyster Crassostrea gigas during the early stages of an outbreak in the Wadden Sea (Germany)

Since the late 1990s, the Pacific oyster (Crassostrea gigas) has spread into the East Frisian Wadden Sea (Germany). This invasion provided an opportunity to study the population dynamics and the patterns of spread during the initial bioinvasion process. With its source area in The Netherlands, the bioinvasion continues in an eastward direction, as documented by a gradient of high abundances in the west and low abundances in the east during the first study year. One year later, abundances of the Pacific oyster were more heterogenic and differed between adjacent tidal basins. The increase in population sizes at all study sites was very high, reaching levels similar to native occurrence populations. The growth constant (K) varied between 0.300 and 0.990 year⁻¹. The mussel bed with the highest densities had a mean abundance of >300 ind. m⁻², and a maximum of 1,460 ind. m⁻². Furthermore, the bioinvasion was facilitated by a low mortality (Z) found for populations between 0.5 and 1.5 years old (Z = 0.03–0.13 year⁻¹). At present, Pacific oysters are well established at several locations in the East Frisian Wadden Sea and may become with these reproductive potential self-sustaining populations.     

Effects of the expansion by Hippophaë rhamnoides on plant species richness in coastal dunes

Question: Is the expansion of Hippophaë rhamnoides in coastal dunes associated with a decline in plant species richness, and is this decline best described by a hump‐backed relationship between species number and shrub cover? Location: Grey and yellow dunes on the East Frisian islands Spiekeroog and Norderney. Methods: Total plant species richness as well as the number of herbaceous and cryptogam species were determined in 2001 using plots of 16 m² size. We compared shrubland plots with varying cover of Hippophaë with neighbouring dune grassland plots without shrubs as reference sites. Soil samples were collected to determine the values of some important edaphic variables (pH, organic matter, nitrogen). Results: The shrubland plots with Hippophaë had or tended to have lower soil pH and C/N ratios and higher contents of organic matter and nitrogen than the grassland plots. Total species richness was marginally significantly related to the cover of Hippophaë in a hump‐backed manner on both islands. The pattern was more pronounced for mosses and lichens than for herbaceous species. For all species groups on Spiekeroog and for the herbaceous species on Norderney, the hump‐backed relationship was much improved when using the difference in species number between shrubland and grassland plot as a dependent variable. Relationships could be improved by including the soil parameters as co‐variables. Species richness was highest at moderate levels of shrub expansion, while it was much reduced in very dense shrubland. The decrease in species number is caused by the decline in grassland species typical of the open dunes, including some rare taxa. Conclusions: The expansion of Hippophaë rhamnoides is a serious threat to the plant species richness of open coastal dunes, and needs to be counteracted by management measures.     

Multiple lines of evidence for demographic and range expansion of a temperate species (Hyla sarda) during the last glaciation

Many temperate species experienced demographic and range contractions in response to climatic changes during Pleistocene glaciations. In this study, we investigate the evolutionary history of the Tyrrhenian tree frog Hyla sarda, a species inhabiting the Corsica-Sardinia island system (Western Mediterranean basin). We used sequence analysis of two mitochondrial (overall 1229 bp) and three nuclear (overall 1692 bp) gene fragments to assess the phylogeography and demographic history of this species, and species distribution modelling (SDM) to predict its range variation over time. Phylogeographic, historical demographic and SDM analyses consistently indicate that H. sarda does not conform to the scenario generally expected for temperate species but rather underwent demographic and range expansion mostly during the last glacial phase. Palaeogeographic data and SDM analyses suggest that such expansion was driven by the glaciation-induced increase in lowland areas during marine regression. This unusual scenario suggests that at least some temperate species may not have suffered the adverse effects of glacial climate on their population size and range extent, owing to the mitigating effects of other glaciations-induced palaeoenvironmental changes. We discuss previous clues for the occurrence of such a scenario in other species and some possible challenges with its identification. Early phylogeographic literature suggested that responses to the Pleistocene glacial-interglacial cycles were expected to vary among species and regions. Our results point out that such variation may have been greater than previously thought.     

A Landscape Perspective of the Stream Corridor Invasion and Habitat Characteristics of an Exotic (Dioscorea oppositifolia) in a Pristine Watershed in Illinois

The spatial distribution of exotics across riparian landscapes is not uniform, and research elaborating the environmental constraints and dispersal behavior that underlie these patterns of distribution is warranted. This study examined the spatial distribution, growth patterns, and habitat constraints of populations of the invasive Dioscorea oppositifolia in a forested stream corridor of a tributary of Drury Creek in Giant City State Park, IL. The distribution of D. oppositifolia was determined at the watershed scale mainly by floodplain structure and connectivity. Populations of D. oppositifolia were confined to the floodplain, with overbank flooding from the stream. Dioscorea oppositifolia probably originates in disturbed areas upstream of natural corridors, and subsequently, the species disperses downstream into pristine canyons or ravines via bulbils dispersing in the water. In Giant City State Park, populations of D. oppositifolia were distributed on the floodplain across broad gradients of soil texture, light, slope, and potential radiation. The study also examined the longevity of bulbils in various micro-environments to illuminate strategies for the management of the species in invaded watersheds. After 1 year, the highest percentages of bulbils were viable under leaves, and much lower percentages were viable over leaves, in soil, and in the creek (76.0±6.8, 21.2±9.6, 21.6±3.6, and 5.2±5.2%), respectively. This study suggests that management procedures that reduce leaf litter on the forest floor (e.g., prescribed burning) could reduce the number of bulbils of D. oppositifolia stored in the watershed.     

Relative contributions of sexual and asexual regeneration strategies in Populus nigra and Salix alba during the first years of establishment on a braided gravel bed river

Populus nigra L. var betulifolia and Salix alba L. var alba are early successional riparian tree species threatened throughout Continental Europe by significant changes to the natural physical processes governing their natural habitat – geomorphologically active floodplains. River management activities have dramatically altered natural patterns of river flow and rates of sediment delivery along rivers, with possible consequences for the balance between sexual and asexual regeneration strategies in these species. Conservation strategies will benefit from a greater understanding of the ways in which dynamic physical processes on the floodplain influence sexual and asexual recruitment. This paper describes a field survey investigating the relative abundance and spatial distributions of P. nigra and S. alba sexual and asexual recruits during the first years of establishment along a braided gravel bed river. Sexual and asexual recruits were identified by excavation along transects in a wet and a dry field season and distributional differences were described in terms of elevation on the floodplain, local sediment type and exposure to floodwaters. Regeneration was overwhelmingly from seed in the first 2–3 years following recruitment, but poor survival rates among sexual recruits saw a shift in the relative abundance of regeneration strategies over time. In relating hydrological data to recruitment, unseasonal flood disturbances had a negative effect on recruitment from seed and a positive effect on vegetative regeneration. Seedlings were associated with fine sediment deposits and were restricted primarily to low elevations on the flood plain, while asexual recruits had a wider spatial distribution. Certain microsite types were unique to either regeneration strategy.     

Use of the green alga Ulva rigida C. Agardh as an indicator species to reassess metal pollution in the Thermaikos Gulf, Greece, after 13 years

The concentrations of metals (Mn, Pb, Fe, Zn, Cu, Cd,Co, Ni, Cr, Na, K, Ca, Mg) were determined in thegreen alga Ulva rigida, in the sediment andseawater of Thermaikos Gulf (Greece) during monthlysamplings in 1994–1995. This Gulf is the recipientof domestic and industrial effluents. Pb, Fe, Cu, Coand Cr concentrations in U. rigida at the studyarea were higher than those 13 years earlier andapparently came from different sources than those forZn, Cd and Ni. The relative abundance of metals inthe alga decreased in the order: Mg > Na > K >Ca > Pb > Fe > Mn > Zn > Cr, Cu > Ni >Co > Cd. Only Cu concentrations in the alga fromKalochori and Cd ones from Viamyl showed significantseasonal changes. Cu and Cd concentrations ingeneral followed the same pattern of variation, withminimum values in winter-spring. This pattern isdiscussed in relation to growth dynamics and tissueage. Only Pb concentrations in the alga showed asignificant positive correlation with concentrationsin the seawater. There were both positive andnegative correlations among some metals in the alga. It is concluded that U. rigida can be used as anindicator species, especially for Pb.     

Downstream changes in the composition,numbers and biomass of bottom fauna in the Tees below Cow Green Reservoir and in an unregulated tributary Maize Beck,in the first five years after impoundment

Changes in composition, numbers and biomass of benthic fauna of the Tees below Cow Green Reservoir and an unregulated tributary Maize Beck were followed between 1972 and 1975 and pre- and post-impoundment conditions were compared. Species diversity was lowest just below the dam and numbers and biomass were highest 240 m downstream of the dam. Faunal densities increased in the Tees after impoundment but in Maize Back no major changes were observed.Freshwater Biological Association, Cow Green UnitFreshwater Biological Association, Cow Green Unit