Genomic admixture increases fitness during a biological invasion

Abstract During biological invasions, multiple introductions can provide opportunities for admixture among genetically distinct lineages. Admixture is predicted to contribute to invasion success by directly increasing fitness through hybrid vigour or by enhancing evolutionary potential within populations . Here, we demonstrate genome‐wide admixture during an invasion that substantially boosted fitness in the cosmopolitan weed, Silene vulgaris. We identified three divergent demes in the native European range that expanded from glacial refugia and experienced historical admixture in a well‐known suture zone. During recent invasion of North America, multiple introductions created additional opportunities for admixture. In common garden experiments, recombinant genotypes from North America experienced a two‐fold increase in fitness relative to nonrecombinants, whereas recombinant genotypes from Europe showed no lasting fitness benefits. This contrast implicates hybrid vigour behind the boost in fitness and supports the hypothesis that admixture can lead to fitness increases that may catapult invasion into a new range.     

Microevolution in Solatopupa landsnails (Pulmonata Chondrinidae): genetic diversity and founder effects

Enzyme variability at 28 presumptive gene loci was studied, by standard starch gel electrophoresis, in 30 populations belonging to the five recognized species of the landsnail Solatopupa from its entire NW Mediterranean range.
Six genetically differentiated groups can be identified among the 30 populations sampled. These are distinguished by two to 19 diagnostic loci, different levels of genetic variability and populations genetics. They are also significantly different as far as the D values are considered. There is no evidence of gene flow among them. Genetically inferred groups correspond in four cases to morphologically distinguished species. In contrast, S. similis , as identified by morphological features, is likely to be a complex of at least two cryptic species.
Populations and species of Solatopupa are characterized by: high fixation of alternative alleles both within and between species; medium-lo-low levels of genie variation; heterozygote deficiency; sharp genetic differentiation among population within species; restricted gene flow; and high genetic distances. Genetic variability is partly associated with climatic factors related to moisture.
Both deterministic and stochastic processes may play a part in the genetic differentiation of Solatopupa snails. Founder events seem to be the main factor affecting the genetic structure of populations and perhaps also speciation. Solatopupa populations display many attributes of populations that may be expected to undergo speciation events involving reorganization of the whole genome after a founder event.     

History, chance and adaptation during biological invasion: separating stochastic phenotypic evolution from response to selection

Introduced species often exhibit changes in genetic variation, population structure, selection regime and phenotypic traits as they colonize and expand into new ranges. For these reasons, species invasions are increasingly recognized as promising systems for studying adaptive evolution over contemporary time scales. However, changes in phenotypic traits during invasion occur under non-equilibrium demographic conditions and may reflect the influences of prior evolutionary history and chance events, as well as selection. We briefly review the evidence for phenotypic evolution and the role of selection during invasion. While there is ample evidence for evolutionary change, it is less clear if selection is the primary mechanism. We then discuss the likelihood that stochastic events shift phenotypic distributions during invasion, and argue that hypotheses of adaptation should be tested against appropriate null models. We suggest two experimental frameworks for separating stochastic evolution from adaptation: statistically accounting for phenotypic variation among putative invasion sources identified by using phylogenetic or assignment methods and by comparing estimates of differentiation within and among ranges for both traits and neutral markers ( Q _ST vs. F _ST). Designs that incorporate a null expectation can reveal the role of history and chance in the evolutionary process, and provide greater insights into evolution during species invasions.     

Plant invasion across space and time: factors affecting nonindigenous species success during four stages of invasion

Invasive nonindigenous plant species (NIPS) threaten native diversity, alter ecosystem processes, and may interact with other components of global environmental change. Here, a general framework is outlined that attempts to connect patterns of plant invasion to processes underlying these patterns at four well-established spatio-temporal stages of the invasion process: transport, colonization, establishment, and landscape spread. At each stage we organize findings and ideas about the filters that limit NIPS success and the interaction of these filters with historical aspects of introduction events, NIPS traits, and ecosystem properties. While it remains difficult to draw conclusions about the risk of invasion across ecosystems, to delineate universal 'invader traits', or to predict large-scale extinctions following invasions, this review highlights the growing body of research that suggests that the success of invasive NIPS is controlled by a series of key processes or filters. These filters are common to all invasion events, and will interact throughout the stages of plant invasion, although the relative importance of a filter may be stage, species or location specific. It is suggested that both research and management programs may benefit from employing multiscale and stage approaches to studying and controlling invasion. We further use the framework to briefly examine potential interactions between climate change and filters that limit NIPS invasion.     

Adaptive divergence and the evolution of reproductive isolation in the wild: an empirical demonstration using introduced sockeye salmon

Populations exposed to different ecological environments should diverge for phenotypic traits that influence survival and reproduction. This adaptive divergence should reduce gene flow between populations because immigrants become less fit than residents and because hybrids perform poorly in either environment (i.e., ecologically-dependent reproductive isolation). Here I demonstrate adaptive divergence and the evolution of reproductive isolation in populations of sockeye salmon (Oncorhynchus nerka) introduced from a common ancestral source into a new lake system (Lake Washington, Washington). The introduced fish founded several new populations, two of which experience very different environments during breeding and early development (Cedar River v.s. Pleasure Point beach). Over 13 generations, the two populations diverged for adult traits (female body size, male body depth; measured in the wild) and embryo traits (survival to hatching, development rate, size at emergence; measured in a common environment). The rates of divergence for these characters were similar to those observed in other examples of rapid evolution, and can best be attributed to natural selection. Partial reproductive isolation has evolved in concert with adaptive divergence: the rate of exchange of adults between the populations (determined using natural tags) is higher than the rate of gene flow (determined using DNA microsatellites). The demonstration that adaptive divergence can initiate reproductive isolation in less than 13 generations suggests that the first signs of ecological speciation may appear soon after new environments are first colonized.     

Genetic Diversity in Introduced Populations with an Allee Effect

A phenomenon that strongly influences the demography of small introduced populations and thereby potentially their genetic diversity is the demographic Allee effect, a reduction in population growth rates at small population sizes. We take a stochastic modeling approach to investigate levels of genetic diversity in populations that successfully overcame either a strong Allee effect, in which populations smaller than a certain critical size are expected to decline, or a weak Allee effect, in which the population growth rate is reduced at small sizes but not negative. Our results indicate that compared to successful populations without an Allee effect, successful populations with a strong Allee effect tend to (1) derive from larger founder population sizes and thus have a higher initial amount of genetic variation, (2) spend fewer generations at small population sizes where genetic drift is particularly strong, and (3) spend more time around the critical population size and thus experience more genetic drift there. In the case of multiple introduction events, there is an additional increase in diversity because Allee-effect populations tend to derive from a larger number of introduction events than other populations. Altogether, a strong Allee effect can either increase or decrease genetic diversity, depending on the average founder population size. By contrast, a weak Allee effect tends to decrease genetic diversity across the entire range of founder population sizes. Finally, we show that it is possible in principle to infer critical population sizes from genetic data, although this would require information from many independently introduced populations.     

Interaction between founder effect and selection during biological invasion in an aquatic plant

Long-distance colonization and rapid range expansion associated with biological invasion may have major evolutionary consequences via both stochastic processes and selection. Using large-scale population genetic surveys, we demonstrate a major shift in the relative frequency of sexually fertile diploid versus sexually sterile triploid populations associated with the invasion of North America by a clonal aquatic plant, Butomus umbellatus. Most populations across the native European range were triploid (84% of 108), whereas most introduced populations were diploid (71% of 136). We evaluated the roles of stochastic processes versus natural selection in causing this shift by surveying predominantly neutral genetic variation at 28 RAPD loci. In Europe (EU) we detected 47 distinct genotypes among 142 plants sampled from 71 populations, whereas in North America (NA) we detected only six genotypes among 138 plants from 69 populations. Of the six NA genotypes, a set of four closely related genotypes were found only in triploid populations and a pair of closely related genotypes were found only in diploid populations, and these were genetically divergent from the triploid genotypes. This result is consistent with severe founder effect. Because sex creates genotypic variation and produces offspring with greater dispersal potential than those produced clonally, we tested the hypothesis that sexual reproduction characteristic of diploids has given them a colonization advantage that accounts for their high frequency in NA. However, we found little or no evidence of sexual recruitment in introduced diploids. One very widespread heterozygous genotype occurred in 95% of 38 introduced diploid populations (i.e., 72 of 76 plants surveyed) suggesting predominant clonal reproduction. Moreover genotypic diversity was not higher within or among diploid than triploid populations in either the native or introduced range. Low genetic diversity in diploid populations was also supported by a comparison of within-population quantitative variation for plant size under a common greenhouse environment. Thus, diploids have not been favored during colonization owing to their sexual fertility. However, concurrent studies have shown that NA diploids exhibit a much higher capacity for clonal reproduction, via small vegetative bulbils, than NA triploids, which almost never produce bulbils. The same difference in clonal capacity is not a consistent feature of the native EU populations. Taken together, these results suggest that strong founder effect has set the stage for a major increase in diploid frequency due to the particular, and possibly idiosyncratic, features of the diploid and triploid lineages introduced to North America.     

Genetic and environmental factors affecting reproductive variation in Allium vineale

Traits related to allocation of resources to sexual and asexual reproduction, together with seed production, were scored on Allium vineale plants sampled from five sites in southern Sweden during a period of 4 years. In addition, random amplified polymorphic DNA (RAPD) fingerprinting of the sampled plants allowed the identification of genets. Integration of genetic and phenotypic data from field and greenhouse provided for the analysis of among‐year, among‐site, and among‐genet variance components. These variance components were taken to represent the influences of short‐term environmental changes, persistent site divergence, and within‐site genet differences, respectively. It was shown that differences among sites and among genets explained a large part of the phenotypic variation of allocation traits, whereas among‐year differences had a larger influence on the variation in seed production. Together, the results support the conclusions of a recent model on the evolution of mixed reproductive systems, that predicts a stable balance between sexual and asexual reproduction because of annual fluctuations in fecundity through the two modes.     

Macroclimatic and maternal effects on the evolution of reproductive traits in lizards

Much of life‐history theory rests on fundamental assumptions about constraints on the acquisition and allocation of energy to growth and reproduction. In general, the allocation of energy to reproduction depends on maternal size, which in turn depends on environmental factors experienced throughout the life of the mother. Here, we used phylogenetic path analyses to evaluate competing hypotheses about the environmental and maternal drivers of reproductive traits in lizards. In doing so, we discovered that precipitation, rather than temperature, has shaped the evolution of the life history. Specifically, environments with greater rainfall have enabled the evolution of larger maternal size. In turn, these larger mothers produce larger clutches of larger offspring. However, annual precipitation has a negative direct effect on offspring size, despite the positive indirect effect mediated by maternal size. Possibly, the evolution of offspring size was driven by the need to conserve water in dry environments, because small organisms are particularly sensitive to water loss. Since we found that body size variation among lizards is related to a combination of climatic factors, mainly precipitation and perhaps primary production, our study challenges previous generalizations (e.g., temperature‐size rule and Bergmann''s rule) and suggests alternative mechanisms underlying the evolution of body size.     

四川大头茶在不同群落中的遗传分化及适合度成分

研究了四川大头茶在 3个群落中的遗传分化和适合度成分。 12个引物的 RAPD分析表明 ,仅有 10 %左右的遗传多样性存在于种群间。四川大头茶纯林结实率 (31.86 % )显著高于针阔混交林 (2 2 .5 3% )和常绿阔叶林 (2 4 .5 2 % )。每果种子数和每果种子均重各种群间差异均显著 ,每果种子数和每果种子均重分别为常绿阔叶林 (2 9.5 0 ,0 .0 15 4 g) ,四川大头茶纯林 (2 8.39,0 .0 172 g) ,针阔混交林 (2 7.4 4 ,0 .0 195 g) ,表明二者间存在着负耦联关系 (trade off)。而以 R =∑lxbx表示的适合度却以常绿阔叶林最大     

Geographic variation and genetic aspects of reproductive diapause in Drosophila triauraria and D. quadraria

ABSTRACT. Geographic variation and genetic aspect of reproductive diapause were studied in Drosophila triauraria Bock & Wheeler and D. quadraria Bock & Wheeler, in relation to their quantitative response to photoperiods. D. quadraria from the subtropical region had no photoperiodic diapause. In D. triauraria strains, diapause was induced under LD 10:14 or 12:12, but not under LD 14:10 or longer photoperiods. Their diapause was induced more effectively and maintained longer by LD 10:14 than by LD 12:12. The duration of diapause was longer in a northern strain, but the diapause inducing process varied little among different geographical strains. Diapause incidence was 50% or lower in F₁ hybrids between D. triauraria and D. quadraria and backcross progenies between F₁ and D. quadraria , and about 70% in backcross progenies between F₁ and D. triauraria under LD 10:14, but very low under LD 12:12. The lower incidence of diapause in these F; and backcross progenies is assumed to be due to the less efficient induction of diapause, since once diapause was induced in them, it was maintained for a long time, especially in the backcross progenies between F₁ and D. triauraria. These experiments suggest that diapause induction and maintenance are different physiological processes controlled by different genetic systems.     

Rapid evolution of Medicago polymorpha during invasion shifts interactions with the soybean looper

The Enemy Release Hypothesis posits that invasion of novel habitats can be facilitated by the absence of coevolved herbivores. However, a new environment and interactions with unfamiliar herbivores may impose selection on invading plants for traits that reduce their attractiveness to herbivores or for enhanced defenses compared to native host plants, leading to a pattern similar to enemy release but driven by evolutionary change rather than ecological differences. The Shifting Defense Hypothesis posits that plants in novel habitats will shift from specialized defense mechanisms to defense mechanisms effective against generalist herbivores in the new range. We tested these ideas by comparing herbivore preference and performance of native (Eurasia)‐ and invasive (New World)‐range Medicago polymorpha, using a generalist herbivore, the soybean looper, that co‐occurs with M. polymorpha in its New World invaded range. We found that soybean loopers varied in preference and performance depending on host genotype and that overall the herbivore preferred to consume plant genotypes from naïve populations from Eurasia. This potentially suggests that range expansion of M. polymorpha into the New World has led to rapid evolution of a variety of traits that have helped multiple populations become established, including those that may allow invasive populations to resist herbivory. Thus, enemy release in a novel range can occur through rapid evolution by the plant during invasion, as predicted by the Shifting Defense Hypothesis, rather than via historical divergence.     

Genetic variation for postzygotic reproductive isolation between Caenorhabditis briggsae and Caenorhabditis sp. 9

The process of speciation is key to the origins of biodiversity, and yet the Caenorhabditis nematode model system has contributed little to this topic. Genetic studies of speciation in the genus are now feasible, owing to crosses between the recently discovered Caenorhabditis sp. 9 and the well-known C. briggsae producing fertile F(1) hybrid females. We dissected patterns of postzygotic reproductive isolation between these species by crossing eight isogenic strains of C. briggsae reciprocally with six strains of C. sp. 9. We determined that overall patterns of reproductive isolation are robust across these genetic backgrounds. However, we also quantified significant heritable variation within each species for interspecific hybrid incompatibilities for total adult progeny, egg-to-adult viability, and the percentage of male progeny. This demonstrates that intraspecific variation for interspecific hybrid incompatibility occurs despite extensive, albeit incomplete, reproductive isolation. Therefore, this emerging general phenomenon of variable reproductive isolation is not restricted to highly interfertile, early-stage incipient species, but also applies to species in the latest stages of the speciation process. Furthermore, we confirm Haldane's rule and demonstrate strongly asymmetric parent-of-origin effects (Darwin's corollary) that consistently manifest more extremely when hermaphroditic C. briggsae serves as maternal parent. These findings highlight Caenorhabditis as an emerging system for understanding the genetics of general patterns of reproductive isolation.     

Repeated and predictable patterns of ecotypic differentiation during a biological invasion: lake–stream divergence in parapatric Swiss stickleback

The relative importance of ecological selection and geographical isolation in promoting and constraining genetic and phenotypic differentiation among populations is not always obvious. Interacting with divergent selection, restricted opportunity for gene flow may in some cases be as much a cause as a consequence of adaptation, with the latter being a hallmark of ecological speciation. Ecological speciation is well studied in parts of the native range of the three‐spined stickleback. Here, we study this process in a recently invaded part of its range. Switzerland was colonized within the past 140 years from at least three different colonization events involving different stickleback lineages. They now occupy diverse habitats, ranging from small streams to the pelagic zone of large lakes. We use replicated systems of parapatric lake and stream populations, some of which trace their origins to different invasive lineages, to ask (i) whether phenotypic divergence occurred among populations inhabiting distinct habitats, (ii) whether trajectories of phenotypic divergence follow predictable parallel patterns and (iii) whether gene flow constrains divergent adaptation or vice versa. We find consistent phenotypic divergence between populations occupying distinct habitats. This involves parallel evolution in several traits with known ecological relevance in independent evolutionary lineages. Adaptive divergence supersedes homogenizing gene flow even at a small spatial scale. We find evidence that adaptive phenotypic divergence places constraints on gene flow over and above that imposed by geographical distance, signalling the early onset of ecological speciation.     

Comparative floral development in male and female plants of Palmer amaranth (Amaranthus palmeri)

Premise

Characterizing the developmental processes in the transition from hermaphroditism to unisexuality is crucial for understanding floral evolution. Amaranthus palmeri, one of the most devastating weeds in the United States, is an emerging model system for studying a dioecious breeding system and understanding the biological traits of this invasive weed. The objectives of this study were to characterize phases of flower development in A. palmeri and compare organogenesis of flower development in female and male plants.

Methods

Flower buds from male and female plants were dissected for light microscopy. Segments of male and female inflorescences at different stages of development were cut longitudinally and visualized using scanning electron microscopy.

Results

Pistillate flowers have two to three styles, one ovary with one ovule, and five obtuse tepals. Staminate flowers have five stamens with five acute tepals. Floral development was classified into 10 stages. The distinction between the two flower types became apparent at stage four by the formation of stamen primordia in staminate flowers, which developed female and male reproductive organs initially, as contrasted to pistillate flowers, which produced carpel primordia only. In staminate flowers, the putative carpel primordia changed little in size and remained undeveloped.

Conclusions

Timing of inappropriate organ termination varies across the two sexes in A. palmeri. Our study suggests that the evolution of A. palmeri from a cosexual ancestral state to complete dioecy is still in progress since males exhibited transient hermaphroditism and females produced strictly pistillate flowers.     

Genetic variation and reproductive patterns in wetland mosses suggest efficient initial colonization of disturbed sites

To understand colonization processes, it is critical to fully assess the role of dispersal in shaping biogeographical patterns at the gene, individual, population, and community levels. We test two alternative hypotheses (H I and H II) for the colonization of disturbed sites by clonal plants, by analyzing intraspecific genetic variation in one and reproductive traits in two typical fen mosses with separate sexes and intermittent spore dispersal, comparing disturbed, early‐succession (limed) fens and late‐successional rich fens. H I suggests initial colonization of disturbed sites by diverse genotypes of which fewer remain in late‐successional fens and an initially balanced sex ratio that develops into a possibly skewed population sex ratio. H II suggests initial colonization by few genotypes and gradual accumulation of additional genotypes and an initially skewed sex ratio that alters into the species‐specific sex ratio, during succession. Under both scenarios, we expect enhanced sexual reproduction in late‐successional fens due to resource gains and decreased intermate distances when clones expand. We show that the intraspecific genetic diversity, assessed by two molecular markers, in Scorpidium cossonii was higher and the genetic variation among sites was smaller in disturbed than late‐successional rich fens. Sex ratio was balanced in S. cossonii and Campylium stellatum in disturbed fens and skewed in C. stellatum in late‐successional fens, thus supporting H I. In line with our prediction, sex expression incidence was higher in, and sporophytes were confined to, late‐succession compared to disturbed rich fens. Late‐successional S. cossonii sites had more within‐site patches with two or more genotypes, and both species displayed higher sex expression levels in late‐successional than in disturbed sites. We conclude that diverse genotypes and both sexes disperse efficiently to, and successfully colonize new sites, while patterns of genetic variation and sexual reproduction in late‐successional rich fens are gradually shaped by local conditions and interactions over extended time periods.     

Artificial introductions, evolutionary change and population differentiation in Trinidadian guppies (Poecilia reticulata; Poeciliidae)

The evolutionary consequences of three artificial introductions of the guppy, Poecilia reticulata , in Trinidad were examined by comparing the allozymic structure (observed heterozygosity ( H _o), and mean number of alleles ( N _a) of each corresponding source (S) and transplant (T) population. In 'Haskins' (H) and EndlerY (E) introduction, 200 guppies (half female) were transferred to guppy-free habitats in 1957 and 1976 respectively. 'Kenny's' (K) introduction in 1981 involved the release of a single pregnant female into an isolated ornamental pond. Analysis of allozyme frequencies at 25 enzyme-coding loci revealed reductions in observed heterozygosity at some loci in all three transplant samples, and a marked decline in the mean number of alleles in Kenny's pond sample. Significant genetic differentiation occurred between (S) and (T) samples at some loci in all introductions, but was most marked in H(T) and K(T). Despite previous studies on rapid evolutionary changes in the life histories and morphology of Endler's transplant guppies, there was little support for any major effects of stochastic forces on allozymic diversity arising from the introduction. Selection arising from changes in predation pressure appeared to be the predominant factor causing the remarkably rapid adaptation of guppies to their new environments. Genetic divergence in some marginal or isolated natural populations was similar to, or greater than, Kenny's pond guppies (Reynolds' genetic distance, R = 0.496), indicating that chance colonization and founder effects may have contributed to the observed geographic patterns of genetic differentiation in Trinidad.     

Mate finding, dispersal, number released, and the success of biological control introductions

Abstract.

• 1 An analysis of published data and a mathematical model of the population dynamics of introduced parasitoids were used to explore the possibility that biological control introductions fail because an Allee effect drives small, introduced populations extinct. Such an Allee effect would arise because low densities, resulting from dispersal into a new environment, lead to failure to mate, which leads to a male-biased sex ratio, which, if extreme enough, could cause population extinction.
• 2 For chalcidoids, ichneumonoids and tachinids, the proportion of parasitoid populations that established when introduced for control of lepidopteran pests increased with the number of parasitoids per release, the total number released, and the number collected when each variable was analysed separately. For chalcidoids alone, establishment increased with the number of releases for this variable analysed separately. However, stepwise logistic regression of establishment on these variables included only the total number released for chalcidoids and the number per release for ichneumonoids and tachinids. This suggests that an Allee effect may limit the establishment of introduced parasitoids more than stochastic environmental variation or lack of genetic variation.
• 3 A reaction-diffusion model of parasitoid introductions was developed, which included mate finding, dispersal, reproduction and survival. Sensitivity analysis showed that the critical number of females needed to establish a population decreased hyperbolically as mate detection distance and net reproductive rate were increased, but the critical number increased linearly as mean-square displacement was increased. The critical number of females did not change when the gross distance traversed per generation was varied. This was because increased area searched by males compensated for increased displacement. Changing from virgin females producing all males (arrhenotoky) to virgin females producing no progeny increased the critical number of females by over 30%.
• 4 The analysis of past introductions and the sensitivity analysis of the reaction-diffusion model both suggested a threshold of about 1000 insects per release to ensure establishment of introduced parasitoids. The implications of our results for the design of biological control introductions are discussed. Limitations in retrospective analyses and current knowledge indicate the need for an experimental approach to introductions.