The role of preadaptation,propagule pressure and competition in the colonization of new habitats

To successfully colonize new habitats, organisms not only need to gain access to it, they also need to cope with the selective pressures imposed by the local biotic and abiotic conditions. The number of immigrants, the preadaptation to the local habitat and the presence of competitors are important factors determining the success of colonization. Here, using two experimental set-ups, we studied the effect of interspecific competition in combination with propagule pressure and preadaptation on the colonization success of new habitats. Our model system consisted of tomato plants (the novel habitat), the two-spotted spider mite Tetranychus urticae as our focal species and the red spider mite Tetranychus evansi as a competitor. Our results show that propagule pressure and preadaptation positively affect colonization success. More successful populations reach larger final population sizes either by having higher per capita growth rates (due to preadaptation effects) or by starting a population with a larger number of individuals. Although populations are more successful colonizing non-competitive environments than competitive ones, propagule pressure and preadaptation counteract the negative effects of competition, promoting colonization success. Our study shows the importance of propagule pressure and preadaptation for successful colonization of new habitats by providing the ability to cope with both the exigencies of new environments and the community context.     

Role of propagule pressure in colonization success: disentangling the relative importance of demographic,genetic and habitat effects

High propagule pressure is arguably the only consistent predictor of colonization success. More individuals enhance colonization success because they aid in overcoming demographic consequences of small population size (e.g. stochasticity and Allee effects). The number of founders can also have direct genetic effects: with fewer individuals, more inbreeding and thus inbreeding depression will occur, whereas more individuals typically harbour greater genetic variation. Thus, the demographic and genetic components of propagule pressure are interrelated, making it difficult to understand which mechanisms are most important in determining colonization success. We experimentally disentangled the demographic and genetic components of propagule pressure by manipulating the number of founders (fewer or more), and genetic background (inbred or outbred) of individuals released in a series of three complementary experiments. We used Bemisia whiteflies and released them onto either their natal host (benign) or a novel host (challenging). Our experiments revealed that having more founding individuals and those individuals being outbred both increased the number of adults produced, but that only genetic background consistently shaped net reproductive rate of experimental populations. Environment was also important and interacted with propagule size to determine the number of adults produced. Quality of the environment interacted also with genetic background to determine establishment success, with a more pronounced effect of inbreeding depression in harsh environments. This interaction did not hold for the net reproductive rate. These data show that the positive effect of propagule pressure on founding success can be driven as much by underlying genetic processes as by demographics. Genetic effects can be immediate and have sizable effects on fitness.     

Biological control as an invasion process: disturbance and propagule pressure affect the invasion success of Lythrum salicaria biological control agents

Understanding the mechanisms behind the successful colonization and establishment of introduced species is important for both preventing the invasion of unwanted species and improving release programs for biological control agents. However, it is often not possible to determine important introduction details, such as date, number of organisms, and introduction location when examining factors affecting invasion success. Here we use biological control introduction data to assess the role of propagule pressure, disturbance, and residence time on invasion success of four herbivorous insect species introduced for the control of the invasive wetland plant, Lythrum salicaria, in the Columbia River Estuary. Two sets of field surveys determined persistence at prior release sites, colonization of new sites, and abundance within colonized sites. We quantified propagule pressure in four ways to examine the effect of different measurements. These included three measurements of introduction size (proximity to introduction site, introduction size at a local scale, and introduction size at a regional scale) and one measure of introduction number (number of introduction events in a region). Disturbance was examined along a tidal inundation gradient (distance from river mouth) and as habitat (island or mainland). Statistical models and model averaging were used to determine which factors were driving invasion success. In this study we found: (1) sparse evidence for the positive influence of propagule pressure on invasion success; (2) disturbance can negatively affect the invasion success of herbivorous insects; (3) the effects of disturbance and propagule pressure are species specific and vary among invasion stages, and (4) not all measures of propagule pressure show the same results, therefore single measures and proxies should be used cautiously.     

Phylogeographic insights into the invasion history and secondary spread of the signal crayfish in Japan

Successful invasion by nonindigenous species is often attributed to high propagule pressure, yet some foreign species become widespread despite showing reduced genetic variation due to founder effects. The signal crayfish (Pacifastacus leniusculus) is one such example, where rapid spread across Japan in recent decades is believed to be the result of only three founding populations. To infer the history and explore the success of this remarkable crayfish invasion, we combined detailed phylogeographical and morphological analyses conducted in both the introduced and native ranges. We sequenced 16S mitochondrial DNA of signal crayfish from across the introduced range in Japan (537 samples, 20 sites) and the native range in western North America (700 samples, 50 sites). Because chela size is often related to aggressive behavior in crayfish, and hence, their invasion success, we also measured chela size of a subset of specimens in both introduced and native ranges. Genetic diversity of introduced signal crayfish populations was as high as that of the dominant phylogeographic group in the native range, suggesting high propagule pressure during invasion. More recently established crayfish populations in Japan that originated through secondary spread from one of the founding populations exhibit reduced genetic diversity relative to older populations, probably as a result of founder effects. However, these newer populations also show larger chela size, consistent with expectations of rapid adaptations or phenotypic responses during the invasion process. Introduced signal crayfish populations in Japan originate from multiple source populations from a wide geographic range in the native range of western North America. A combination of high genetic diversity, especially for older populations in the invasive range, and rapid adaptation to colonization, manifested as larger chela in recent invasions, likely contribute to invasion success of signal crayfish in Japan.     

Allozyme variability in an invasive drosophilid,Zaprionus indianus (Diptera: Drosophilidae): comparison of a recently introduced Brazilian population with Old World populations

Colonizing species often go through genetic bottlenecks when new territories are invaded. The South American continent has been recently colonized by a generalist African drosophilid, Zaprionus indianus, which has become an agricultural pest in Brazil in the last five years. In this paper we used allozyme electrophoresis to estimate levels of genetic differentiation of Z. indianus collected from sites 4 300 km apart in Brazil. We also compared the level of polymorphism of the Brazilian populations with that found in laboratory strains from Africa and Asia, to verify if a significant decrease in gene variability has taken place during the invasion process. The populations were polymorphic for three out of the 11 loci investigated. Genetic distances and F_ST indices among Brazilian populations were small and generally non significant, suggesting a colonization from one single propagule followed by a rapid demographic expansion. Ancestral and old populations from Africa and Asia were slightly more heterozygous than those from Brazil. Compared to other drosophilids, Z. indianus appears to be characterized by a low proportion (25%) of polymorphic loci. We suggest that the propagule introduced to Brazil had a sufficient size to carry almost all the polymorphism from the (unknown) origin population, although not the precise allelic frequencies.     

Parsing propagule pressure: Number,not size,of introductions drives colonization success in a novel environment

Predicting whether individuals will colonize a novel habitat is of fundamental ecological interest and is crucial to conservation efforts. A consistently supported predictor of colonization success is the number of individuals introduced, also called propagule pressure. Propagule pressure increases with the number of introductions and the number of individuals per introduction (the size of the introduction), but it is unresolved which process is a stronger driver of colonization success. Furthermore, their relative importance may depend upon the environment, with multiple introductions potentially enhancing colonization of fluctuating environments. To evaluate the relative importance of the number and size of introductions and its dependence upon environmental variability, we paired demographic simulations with a microcosm experiment. Using Tribolium flour beetles as a model system, we introduced a fixed number of individuals into replicated novel habitats of stable or fluctuating quality, varying the number of introductions through time and size of each introduction. We evaluated establishment probability and the size of extant populations through seven generations. We found that establishment probability generally increased with more, smaller introductions, but was not affected by biologically realistic fluctuations in environmental quality. Population size was not significantly affected by environmental variability in the simulations, but populations in the microcosms grew larger in a stable environment, especially with more introduction events. In general, the microcosm experiment yielded higher establishment probability and larger populations than the demographic simulations. We suggest that genetic mechanisms likely underlie these differences and thus deserve more attention in efforts to parse propagule pressure. Our results highlight the importance of preventing further introductions of undesirable species to invaded sites and suggest conservation efforts should focus on increasing the number of introductions or reintroductions of desirable species rather than increasing the size of those introduction events into harsh environments.     

FITNESS CONSEQUENCES OF OUTCROSSING IN A SOCIAL SPIDER WITH AN INBREEDING MATING SYSTEM

Inbreeding mating systems are uncommon because of inbreeding depression. Mating among close relatives can evolve, however, when outcrossing is constrained. Social spiders show obligatory mating among siblings. In combination with a female‐biased sex ratio, sib‐mating results in small effective populations. In such a system, high genetic homozygosity is expected, and drift may cause population divergence. We tested the effect of outcrossing in the social spider Stegodyphus dumicola. Females were mated to sib‐males, to a non‐nestmate within the population, or to a male from a distant population, and fitness traits of F1s were compared. We found reduced hatching success of broods from between‐population crosses, suggesting the presence of population divergence at a large geographical scale that may result in population incompatibility. However, a lack of a difference in offspring performance between inbred and outbred crosses indicates little genetic variation between populations, and could suggest recent colonization by a common ancestor. This is consistent with population dynamics of frequent colonizations by single sib‐mated females of common origin, and extinctions of populations after few generations. Although drift or single mutations can lead to population divergence at a relatively short time scale, it is possible that dynamic population processes homogenize these effects at longer time scales.     

How propagule size and environmental suitability jointly determine establishment success: a test using dung beetle introductions

Both the size of founding populations (propagule size) and environmental suitability are known to influence whether a species newly introduced to a location will establish a self-sustaining population. However, these two factors do not operate independently: it is the interaction between propagule size and environmental suitability that determines the probability an introduced population will establish. Here I use the example of dung beetle introductions to Australia to illustrate the importance of this interaction. I first describe equations that model establishment success jointly as a function of propagule size and environmental suitability. I then show how these equations provide insight into the different outcomes observed in two dung beetle species widely introduced to Australia. In one species, variation in propagule size had relatively little influence on establishment success due to large variation in environmental suitability, leading to an essentially bimodal outcome: sites were either very suitable for establishment and introductions succeeded, or sites were unsuitable and introductions failed regardless of propagule size. For the second species, there was much less variation among locations in environmental suitability, leading to propagule size having a strong influence on establishment success. These examples highlight how the interplay between environmental suitability and founding population size is central to determining the probability an introduced species will establish.     

Population characteristics of Gammarus pulex (L.) from five English streams

Gammarus pulex were sampled from five English streams during April 1992. The population density, number of precopula pairs and incidence of parasitic infection were recorded, and the biomass was estimated from subsamples by relating body area to dry weight. Physical and chemical measurements were taken from each stream. The abundance and standing crop biomass differed significantly between streams, probably due to the influence of pollutants or the physical structure of the stream bed. The size of individual G. pulex also differed significantly between streams, although there was no obvious causal explanation for this. Few individuals were visibly parasitised in any of the populations. Males were significantly larger than females, both in precopula pairs and in the general populations. The sex ratio differed between populations and may explain inter-stream differences in the relationship between precopula male and female size.     

Prior adaptation,diversity, and introduction frequency mediate the positive relationship between propagule pressure and the initial success of founding populations

Propagule pressure is often considered the most consistent predictor of the success of founding populations. This relationship could be mediated by the composition of the founding group (e.g. level of prior adaptation to the recipient environment or its diversity) as well as the introduction scenario (i.e. the frequency, size and timing of discrete introduction events). We introduced groups of Tribolium castaneum (red flour beetle) eggs across three levels of propagule pressure (n?=?15, 30, 60), of three possible compositions (single, adapted lineage; single, unadapted lineage; mixed lineages) to a novel environment using six unique introduction scenarios, in a fully factorial design to evaluate the importance of composition and introduction scenario in influencing the relationship between propagule pressure and establishment. In our system, prior adaptation to the environment, including having some adapted individuals in mixed groups, rivaled the importance of propagule pressure in determining the establishment success and size of founding populations. More frequent introduction events resulted in fewer individuals that initially survived founding, but introduction scenario did not significantly influence establishment success or population size. This experimental evidence demonstrates the importance of context, both of the founding group and the recipient environment, in understanding how propagule pressure influences the success of founding populations.     

Accentuated polymorphism of heterochromatin and nucleolar organizer regions in Astyanax scabripinnis (Pisces, Characidae): tools for understanding karyotypic evolution

Astyanax scabripinnis has been considered a species complex because it presents high karyotypic and morphological variability among its populations. In this work, individuals of two A. scabripinnis populations from different streams in the same hydrographic basin were analyzed through C‐banding and AgNOR. Although they present distinct diploid numbers, they show meta and submetacentric chromosome groups highly conserved (numerically and morphologically). Other chromosomal characteristics are also shared by both populations, as the pattern of constitutive heterochromatin distribution (large blocks in the telomeric regions of subtelocentric and acrocentric chromosomes) and some nucleolar chromosomes. Inter‐individual variations both in the number and size of heterochromatic blocks, and in the number and localization of NORs were verified in the studied populations, characterizing them as polymorphics for these regions. The mechanisms involved in the dispersion of heterochromatin and NORs through the karyotypes, as well as the possible events related to the generation of polymorphism of those regions are discussed. Furthermore, relationships between these populations and within the context of the scabripinnis complex are also approached. This revised version was published online in July 2006 with corrections to the Cover Date.     

Genetic signals of artificial and natural dispersal linked to colonization of South America by non‐native Chinook salmon (Oncorhynchus tshawytscha)

Genetics data have provided unprecedented insights into evolutionary aspects of colonization by non‐native populations. Yet, our understanding of how artificial (human‐mediated) and natural dispersal pathways of non‐native individuals influence genetic metrics, evolution of genetic structure, and admixture remains elusive. We capitalize on the widespread colonization of Chinook salmon Oncorhynchus tshawytscha in South America, mediated by both dispersal pathways, to address these issues using data from a panel of polymorphic SNPs. First, genetic diversity and the number of effective breeders (N_b) were higher among artificial than natural populations. Contemporary gene flow was common between adjacent artificial and natural and adjacent natural populations, but uncommon between geographically distant populations. Second, genetic structure revealed four distinct clusters throughout the Chinook salmon distributional range with varying levels of genetic connectivity. Isolation by distance resulted from weak differentiation between adjacent artificial and natural and between natural populations, with strong differentiation between distant Pacific Ocean and Atlantic Ocean populations, which experienced strong genetic drift. Third, genetic mixture analyses revealed the presence of at least six donor geographic regions from North America, some of which likely hybridized as a result of multiple introductions. Relative propagule pressure or the proportion of Chinook salmon propagules introduced from various geographic regions according to government records significantly influenced genetic mixtures for two of three artificial populations. Our findings support a model of colonization in which high‐diversity artificial populations established first; some of these populations exhibited significant admixture resulting from propagule pressure. Low‐diversity natural populations were likely subsequently founded from a reduced number of individuals.     

Forest fragmentation and plant reproductive success: a case study in four perennial herbs

We assessed the effects of habitat fragmentation on reproductive success in natural populations of four forest herbs with differing life-history traits and whose distribution patterns appeared to be negatively affected by decreased habitat size and/or increased isolation: Carex sylvatica, Galium odoratum, Sanicula europaea and Veronica montana. Our aims were to test (1) whether habitat size and isolation are positively correlated with population size and isolation, respectively, (2) whether plant reproductive success, a major component of plant fitness, is reduced in small and/or isolated populations when also accounting for differences in habitat quality (edaphic conditions, light intensity) and the effects of plant size, and (3) whether species with different life histories are affected differently. There were significant positive relationships between habitat and population size and between habitat and population isolation in some, but not all of the species. We mostly found no negative effects of small population size or isolation on reproduction. However, reproductive success was reduced in small populations of Sanicula, and this effect was independent of differences in plant size and environmental conditions. The reduced fecundity in small populations may be a consequence of the Allee-effect, a possible mechanism being pollen limitation. Furthermore, the proportion of flowering ramets was reduced in small and isolated populations of Galium, which may have been caused by changes in population structure. Lastly, we found some evidence for largely outcrossing, non-clonal species to be more sensitive to reductions in population size, at least in terms of their reproductive success.     

Population‐level traits that affect,and do not affect,invasion success

What allows some species to successfully colonize a novel environment while others fail? Numerous studies in invasion biology have sought to answer this question, but those studies have tended to focus on traits of species or individuals (e.g. body size, seed size, seed number), and these traits have largely been found to be weak predictors of invasion success. However, characteristics of colonizing populations (e.g. genetic diversity, density, age structure) might also be important for successful establishment, as the authors of a study published in this issue of Molecular Ecology show ( Crawford & Whitney 2010 ). By experimentally manipulating the density and genetic diversity of colonizing populations of Arabidopsis thaliana, the authors found that genetic diversity, but not population density, increased colonization success. Importantly, the effects of genetic diversity on colonization success were both additive and non‐additive, suggesting that traits associated with particular genotypes and complimentarity among genotypes contribute to colonization success. This research highlights the importance of considering within‐species variation and characteristics of entire populations in predicting colonization success.     

Aquaculture facilities drive the introduction and establishment of non-native Oreochromis niloticus populations in Neotropical streams

This study investigated whether aquaculture facilities drive the introduction and establishment of the non-native Nile tilapia (Orechromis niloticus) in Neotropical streams, Brazil. Samples were taken from nine streams with different aquaculture occupation intensities (no, moderate, and intense) using the percentage of occupation of micro-watersheds by aquaculture ponds as a proxy for propagule pressure. The presence of aquaculture facilities and the percentage of aquaculture occupation were good predictors of the catch frequency and of densities of tilapias in the adjacent natural environment. In the streams under intense propagule pressure, females prevailed and high densities of young individuals comparing to adults were recorded. It suggests that the species was reproductively successful in the natural environment. In the streams under moderate propagule pressure, males prevailed, which indicates the likely capture of individuals escaped from aquaculture facilities. In general, our results show the positive influence of propagule pressure on the introduction and establishment of Nile tilapia in natural ecosystems, showing that aquaculture expansion of O. niloticus poses a threat to the conservation of aquatic biodiversity.

     

The relative importance of latitude matching and propagule pressure in the colonization success of an invasive forb

Factors that influence the early stages of invasion can be critical to invasion success, yet are seldom studied. In particular, broad pre-adaptation to recipient climate may importantly influence early colonization success, yet few studies have explicitly examined this. I performed an experiment to determine how similarity between seed source and transplant site latitude, as a general indicator of pre-adaptation to climate, interacts with propagule pressure (100, 200 and 400 seeds/pot) to influence early colonization success of the widespread North American weed, St. John's wort Hypericum perforatum . Seeds originating from seven native European source populations were sown in pots buried in the ground in a field in western Montana. Seed source populations were either similar or divergent in latitude to the recipient transplant site. Across seed density treatments, the match between seed source and recipient latitude did not affect the proportion of pots colonized or the number of individual colonists per pot. In contrast, propagule pressure had a significant and positive effect on colonization. These results suggest that propagules from many climatically divergent source populations can be viable invaders.     

Liriomyza huidobrensis in Yunnan, China: current distribution and genetic structure of a recently established population

Liriomyza huidobrensis Blanchard (Diptera: Agromyzidae) is a very serious and economically important pest around the world. Liriomyza huidobrensis in China was first reported from Kunming of Yunnan province in 1993. We report here that this pest has recently expanded its distribution, along with a host plant range extension and population explosion. The mitochondrial cytochrome oxidase II gene was sequenced for eight populations from Yunnan. All individuals were identical: no genetic variation was observed between populations either from different geographical localities or from different host plants. The phylogenetic analysis shows that the Yunnan population is grouped into the South American clade, which also includes other recently introduced Asian populations. Together with ecological data and colonization history of this pest, our results suggest that Yunnan population might have an ultimate, albeit not immediate, origin from South American populations.     

Experimental studies of adaptive differentiation in Bahamian Anolis lizards

Populations of the lizards Anolis carolinensis and A. sagrei were experimentally introduced onto small islands in the Bahamas. Less than 15 years after introduction, we investigated whether the populations had diverged and, if so, whether differentiation was related to island vegetational characteristics or propagule size. No effect of founding population size was evident, but differentiation of A. sagrei appears to have been adaptive, a direct relationship existed between how vegetationally different an experimental island was from the source island and how much the experimental population on that island had diverged morphologically. Populations of A. carolinensis had also diverged, but were too few for quantitative comparisons. A parallel exists between the divergence of experimental populations of A. sagrei and the adaptive radiation of Anolis lizards in the Greater Antilles; in both cases, relative hindlimb length and perch diameter are strongly correlated. This differentiation could have resulted from genetic change or environmentally-driven phenotypic plasticity. Laboratory studies on A. sagrei from a population in Florida indicate that hindlimb length exhibits adaptive phenotypic plasticity. Further studies are required to determine if the observed differences among the experimental populations are the result of such plasticity. Regardless of whether the differences result from plasticity, genetic change, or both, the observation that anole populations differentiate rapidly and adaptively when exposed to novel environmental conditions has important implications for understanding the adaptive radiation of Caribbean anoles.     

Determinants of vertebrate invasion success in Europe and North America

Species that are frequently introduced to an exotic range have a high potential of becoming invasive. Besides propagule pressure, however, no other generally strong determinant of invasion success is known. Although evidence has accumulated that human affiliates (domesticates, pets, human commensals) also have high invasion success, existing studies do not distinguish whether this success can be completely explained by or is partly independent of propagule pressure. Here, we analyze both factors independently, propagule pressure and human affiliation. We also consider a third factor directly related to humans, hunting, and 17 traits on each species' population size and extent, diet, body size, and life history. Our dataset includes all 2362 freshwater fish, mammals, and birds native to Europe or North America. In contrast to most previous studies, we look at the complete invasion process consisting of (1) introduction, (2) establishment, and (3) spread. In this way, we not only consider which of the introduced species became invasive but also which species were introduced. Of the 20 factors tested, propagule pressure and human affiliation were the two strongest determinants of invasion success across all taxa and steps. This was true for multivariate analyses that account for intercorrelations among variables as well as univariate analyses, suggesting that human affiliation influenced invasion success independently of propagule pressure. Some factors affected the different steps of the invasion process antagonistically. For example, game species were much more likely to be introduced to an exotic continent than nonhunted species but tended to be less likely to establish themselves and spread. Such antagonistic effects show the importance of considering the complete invasion process.