Invasive species are predicted to suffer from reductions in genetic diversity during founding events, reducing adaptive potential. Integrating evidence from two literature reviews and two case studies, we address the following questions: How much genetic diversity is lost in invasions? Do multiple introductions ameliorate this loss? Is there evidence for loss of diversity in quantitative traits? Do invaders that have experienced strong bottlenecks show adaptive evolution? How do multiple introductions influence adaptation on a landscape scale? We reviewed studies of 80 species of animals, plants, and fungi that quantified nuclear molecular diversity within introduced and source populations. Overall, there were significant losses of both allelic richness and heterozygosity in introduced populations, and large gains in diversity were rare. Evidence for multiple introductions was associated with increased diversity, and allelic variation appeared to increase over long timescales (~100 years), suggesting a role for gene flow in augmenting diversity over the long-term. We then reviewed the literature on quantitative trait diversity and found that broad-sense variation rarely declines in introductions, but direct comparisons of additive variance were lacking. Our studies of Hypericum canariense invasions illustrate how populations with diminished diversity may still evolve rapidly. Given the prevalence of genetic bottlenecks in successful invading populations and the potential for adaptive evolution in quantitative traits, we suggest that the disadvantages associated with founding events may have been overstated. However, our work on the successful invader Verbascum thapsus illustrates how multiple introductions may take time to commingle, instead persisting as a 'mosaic of maladaptation' where traits are not distributed in a pattern consistent with adaptation. We conclude that management limiting gene flow among introduced populations may reduce adaptive potential but is unlikely to prevent expansion or the evolution of novel invasive behaviour.

/pdf/founding-events-in-species-invasions-genetic-variation-138f2ii5cu.pdf

Founding events in species invasions: genetic variation, adaptive evolution, and the role of multiple introductions

LEARNING The objectives of BIOS 781 are to present: OBJECTIVES: 1. basic population and quantitative genetic principles, including classical genetics, chromosomal theory of inheritance, and meiotic recombination 2. an exposure to QTL mapping methods of complex quantitative traits and linkage methods to detect co-segregation with disease 3. methods for assessing marker-disease linkage disequilibrium, including case-control approaches 4. methods for genome-wide association and stratification control.

Genetic Data Analysis

Invasions by non-native ants are an ecologically destructive phenomenon affecting both continental and island ecosystems throughout the world. Invasive ants often become highly abundant in their introduced range and can outnumber native ants. These numerical disparities underlie the competitive asymmetry between invasive ants and native ants and result from a complex interplay of behavioral, ecological, and genetic factors. Reductions in the diversity and abundance of native ants resulting from ant invasions give rise to a variety of direct and indirect effects on non-ant taxa. Invasive ants compete with and prey upon a diversity of other organisms, including some vertebrates, and may enter into or disrupt mutualistic interactions with numerous plants and other insects. Experimental studies and research focused on the native range ecology of invasive ants will be especially valuable contributions to this field of study.

The Causes and Consequences of Ant Invasions

The phenomenon of inbreeding depression is well documented and behavioral adaptations for inbreeding avoidance have been described. However, there is debate over whether inbreeding depression is always an important selective force on behavior. Here, we summarize recent evidence for inbreeding depression under natural conditions, review inbreeding avoidance mechanisms, and discuss how these are influenced by social structure. We also examine the idea that animals have evolved mechanisms to avoid outbreeding.

Inbreeding avoidance in animals.

Despite the severe ecological and economic damage caused by introduced species, factors that allow invaders to become successful often remain elusive. Of invasive taxa, ants are among the most widespread and harmful. Highly invasive ants are often unicolonial, forming supercolonies in which workers and queens mix freely among physically separate nests. By reducing costs associated with territoriality, unicolonial species can attain high worker densities, allowing them to achieve interspecific dominance. Here we examine the behavior and population genetics of the invasive Argentine ant (Linepithema humile) in its native and introduced ranges, and we provide a mechanism to explain its success as an invader. Using microsatellite markers, we show that a population bottleneck has reduced the genetic diversity of introduced populations. This loss is associated with reduced intraspecific aggression among spatially separate nests, and leads to the formation of interspecifically dominant supercolonies. In contrast, native populations are more genetically variable and exhibit pronounced intraspecific aggression. Although reductions in genetic diversity are generally considered detrimental, these findings provide an example of how a genetic bottleneck can lead to widespread ecological success. In addition, these results provide insights into the origin and evolution of unicoloniality, which is often considered a challenge to kin selection theory.

/pdf/reduced-genetic-variation-and-the-success-of-an-invasive-15mkcip8ye.pdf

Reduced genetic variation and the success of an invasive species

/pdf/a-revision-of-the-fire-ants-solenopsis-geminata-group-2o6by2zhb2.pdf

A revision of the fire ants, Solenopsis geminata group (Hymenoptera: Formicidae, Myrmicinae).

Effects of a recent founder event on genetic diversity in wild populations of the fire ant Solenopsis invicta were studied, with particular attention given to the genetic sex-determining system. Diploid males are far more common relative to haploid males in introduced populations than in native populations of fire ants, and queens that produce diploid males account for a significantly larger proportion of the mated queens in introduced than in native populations. Differences between native and introduced populations in attributes of the mating systems (i.e., queen mating frequency or level of inbreeding) can be excluded as factors contributing to these different levels of diploid male production. Thus, we conclude that diploid males have increased in frequency in introduced populations because of a loss of allelic diversity at the sex-determining locus (loci). This loss of sex alleles has generated a substantial increase in the estimated segregational genetic load associated with production of sterile diploid males in introduced populations over the load in native populations. The loss of allelic diversity in the sex-determining system in introduced S. invicta is paralleled by a loss of electrophoretically detectable rare alleles at protein-encoding loci. Such concordance between these different types of markers is predicted because each of the many sex alleles present in the native populations is expected to be rare. Estimates of expected heterozygosity (Hexp) based on 76 electrophoretic loci do not differ significantly between the native and introduced fire ant populations, illustrating the lack of sensitivity of this measure for detecting many types of bottlenecks.

https://www.genetics.org/content/genetics/135/3/843.full.pdf

Effect of a founder event on variation in the genetic sex-determining system of the fire ant Solenopsis invicta.

A new survey of the ants of the Florida Keys increases the known fauna from 30 to 83 species. An annotated list provides data on habitats, collection sites, and location of vouchers. Solenopsis corticalis Forel, Leptothorax torrei (Aguayo) and Monomorium ebeninum Forel are new records for the U.S. The fauna includes 27 exotics and 31 species native to the southeastern Coastal Plain; most of the remaining species are Antillean. There are 2 possibly endemic species. The proportion of known exotics (33%) in the fauna is the highest for any area in the U.S. There is evidence that populations of certain exotics are increasing with increasing disturbance of remaining native habitats. Species diversity in the Keys is probably limited by 1) limited habitat diversity, 2) lack of easy access to the rich ant fauna of the Neotropics, and 3) unsuitability of the climate and habitats for many species found farther north.

A review of the ants of the Florida Keys

Many species of plants and animals associated with grasslands are rare or declining due to habitat loss and degradation. Although grassland plants and insects evolved in the context of both grazing and fire, the appropriate use of grazing and fire has been debated among those concerned with protecting insect communities. We established an experiment to test insect responses to three grassland management treatments: (1) patch-burn graze (burning of spatially distinct patches and free access by cattle), (2) graze- and-burn (burning of entire tract with free access by cattle), and (3) burn-only. Because we expected that land-use legacies could also affect insect abundance and diversity, we evaluated effects of time since fire, grazing history, remnant history (remnant or reconstructed grassland) and pre-treatment vegetation characteristics, which were assumed to be a legacy of prior land-use. Butterflies (Lepidoptera), ants (Hymenoptera: Formicidae), and leaf beetles (Coleoptera: Chrysomelidae) were surveyed for three years to compare their responses to each of these treatments as measured by abundance, richness and species diversity. Each of these taxa is relatively diverse and was expected to have the potential to have strong negative responses to grazing and burning, but we predicted more positive responses to patch-burn grazing. Our results showed that land-use legacies affected insect abundance, richness and diversity, but treatments did not. Ant abundance was lower in tracts with a history of heavy grazing. Ant species richness was positively associated with pre-treatment time since fire and vegetation height and negatively associated with pre-treatment proportion native plant cover. Butterfly abundance was positively associated with pre-treatment litter cover. Leaf beetle diversity was positively associated with pre-treatment native plant cover, and leaf beetle abundance was negatively associated with time since fire. Our results indicate that land-use legacies can exert more influence on grassland insect community composition than current management, but the particular aspects of these land-use legacies that are important vary across insect taxa. The implications of these finding are that (1) land-use legacies should garner more attention in grassland management and (2) conservation of grassland insect communities will be improved by taxon- specific analysis of land-use legacy variables.

http://millerlab.nres.illinois.edu/pdfs/Debinski_2011_Cross-taxonomic%20comparison%20of%20insect%20responses%20to%20grassland%20management.pdf

A Cross-Taxonomic Comparison of Insect Responses to Grassland Management and Land-Use Legacies

Specimens of seven fire ant species collected from their native ranges in Argentina were studied by protein electrophoresis and morphological analysis. Concordance between the genetic and morphological character sets is strong (96% agreement on identifications), suggesting that recognition of reproductively isolated populations and partitioning of intra- and interspecific variation can in most cases be achieved using appropriate characters of either type in this taxonomically difficult group. Genetic differentiation between native (Argentina) and introduced (USA) conspecific populations of two species, Solenopsis invicta and S. richteri, is rather typical of the differentiation existing between conspecific populations found within either country. Furthermore, there appears to have been little reduction of variability (heterozygosity) at enzyme loci following colonization by either species of the United States, although some rare alleles have been lost in the introduced populations. Hybridization is rare between S. invicta and S. richteri where their native ranges overlap in central Argentina, in contrast to the extensive hybridization of these species in the United States, suggesting that prezygotic barriers to gene flow have been compromised in introduced populations. Phylogenetic analysis of the seven species indicates that S. invicta and S. richteri are relatively distantly related within the S. saevissima complex. Given that hybrids between these species in the United States suffer little apparent loss of fitness, genomic incompatibilities generally may be insufficient to create effective postzygotic barriers to interspecific gene flow in this group of ants.

https://onlinelibrary.wiley.com/doi/pdf/10.1111/j.1558-5646.1990.tb04316.x

James C. Trager

Papers

A revision of the fire ants, Solenopsis geminata group (Hymenoptera: Formicidae, Myrmicinae).

Effect of a founder event on variation in the genetic sex-determining system of the fire ant Solenopsis invicta.

A review of the ants of the Florida Keys

A Cross-Taxonomic Comparison of Insect Responses to Grassland Management and Land-Use Legacies

Systematics and population genetics of fire ants (solenopsis saevissima complex) from argentina.