Variation in the level of aggression, chemical and genetic distance among three supercolonies of the Argentine ant in Europe

In their invasive ranges, Argentine ant populations often form one geographically vast supercolony, genetically and chemically uniform within which there is no intraspecific aggression. Here we present regional patterns of intraspecific aggression, cuticular hydrocarbons (CHCs) and population genetics of 18 nesting sites across Corsica and the French mainland. Aggression tests confirm the presence of a third European supercolony, the Corsican supercolony, which exhibits moderate to high levels of aggression, depending on nesting sites, with the Main supercolony, and invariably high levels of aggression with the Catalonian supercolony. The chemical analyses corroborated the behavioural data, with workers of the Corsican supercolony showing moderate differences in CHCs compared to workers of the European Main supercolony and strong differences compared to workers of the Catalonian supercolony. Interestingly, there were also clear genetic differences between workers of the Catalonian supercolony and the two other supercolonies at both nuclear and mitochondrial markers, but only very weak genetic differentiation between nesting sites of the Corsican and Main supercolonies (F(ST) = 0.06). A detailed comparison of the genetic composition of supercolonies also revealed that, if one of the last two supercolonies derived from the other, it is the Main supercolony that derived from the Corsican supercolony rather than the reverse. Overall, these findings highlight the importance of conducting more qualitative and quantitative analyses of the level of aggression between supercolonies, which has to be correlated with genetic and chemical data.     

外来种入侵的生物学与生态学基础的若干问题

外来物种的入侵及其所造成的危害已越来越为人们所认识。一个外来物种入侵一个新的生境无疑要具备一定的条件 ,尽管目前对这种条件的研究还只是初步的。首先从其自身来说 ,需要有足够的入侵性 ,这可以认为是入侵种的生物学基础 ,可能表现为具有多倍性、一定的遗传变异、杂合性或表型可塑性 ,以及不同的交配系统等 ;其次一个物种的入侵成功还取决于入侵生境的可入侵性 ,这可以认为是入侵种的生态学基础 ,而新的种间关系的形成 ,尤其是入侵种与其他种类的互惠共生关系的建成是关键。本文主要介绍了有关入侵种的生物学和生态学基础中的一些最新进展 ,希望有助于我国学者对相关研究内容的开展以及对有害外来种入侵的防治     

The role of cuticular hydrocarbons as chemical cues for nestmate recognition in the invasive Argentine ant (<Emphasis Type="Italic">Linepithema humile</Emphasis>)

Argentine ants (Linepithema humile) in their native South American range, like most other ant species, form spatially restricted colonies that display high levels of aggression toward other such colonies. In their introduced range, Argentine ants are unicolonial and form massive supercolonies composed of numerous nests among which territorial boundaries are absent. Here we examine the role of cuticular hydrocarbons (CHCs) in nestmate recognition of this highly damaging invasive ant using three supercolonies from its introduced range. We conducted behavioral assays to test the response of Argentine ants to workers treated with colonymate or non-colonymate CHCs. Additionally, we quantified the amount of hydrocarbons transferred to individual ants and performed gas chromatography-mass spectrometry (GC/MS) to qualitatively characterize our manipulation of CHC profiles. The GC/MS data revealed marked differences in the hydrocarbon profiles across supercolonies and indicated that our treatment effectively masked the original chemical profile of the treated ants with the profile belonging to the foreign individuals. We found that individual workers treated with foreign CHCs were aggressively rejected by their colonymates and this behavior appears to be concentration-dependent: larger quantities of foreign CHCs triggered higher levels of aggression. Moreover, this response was not simply due to an increase in the amount of CHCs applied to the cuticle since treatment with high concentrations of nestmate CHCs did not trigger aggression.The results of this study bolster the findings of previous studies on social insects that have implicated CHCs as nestmate recognition cues and provide insight into the mechanisms of nestmate recognition in the invasive Argentine ant. Received 6 February 2007; revised 31 May and 27 July 2007; accepted 16 August 2007.     

Molecular analysis of the efficiency of sloughed skin sampling in whale population genetics

Sloughed whale skin contains enough DNA for genetic analysis, and offers a nonintrusive method for collecting tissue. Here, we examine the efficiency of sloughed skin sampling using 1460 samples collected from free-ranging humpback whales. Samples were sexed and screened for up to 10 microsatellite markers. The vast majority of samples appear genetically compatible with field observations. About 1% of groups revealed more genotypes than whales, but we argue that this is more likely to be due to unobserved whales than to contamination. Sloughed skin sampling is particularly effective when applied to active groups and offers a viable alternative to biopsy darting in regions where darting is either not permitted or otherwise undesirable.     

The diminutive supercolony: the Argentine ants of the southeastern United States

Native to Argentina and Brazil, the Argentine ant (Linepithema humile) is an invasive species that has become established on six continents and many oceanic islands. In several parts of its introduced range, including the western United States, southern Europe and Chile, the Argentine ant is unicolonial, forming extensive supercolonies. We examined population genetic structure and intercolony aggression in two regions of the introduced range of this species in the United States: California and the southeastern United States. Our results show that the southeastern L. humile population has high genotypic variability and strong intercolony aggression relative to the California population. In the California population, intercolony aggression was absent and 23 alleles were found across seven polymorphic microsatellite loci. However, in the Southeast, aggression between colonies was high and 47 alleles were present across the same seven loci in an equal number of colonies. We suggest that distinctly different colonization patterns for California and the Southeast may be responsible for the striking disparity in the genetic diversity of introduced populations. Southeastern colonies may have descended from multiple, independent introductions from the native range, undergoing a bottleneck at each introduction. In contrast, the California supercolony may have originated from one or more colonies inhabiting the southeastern United States, thus experiencing a double bottleneck. The differences in present-day distribution patterns between California and the Southeast may be due to the combined effect of two factors: lower winter temperatures in the Southeast and/or competition with another successful and widely distributed ant invader, the fire ant Solenopsis invicta.     

Isolation and characterization of microsatellite loci from the invasive ant Pheidole megacephala

We report the characterization of eight microsatellite markers in the big-headed ant Pheidole megacephala, a pest ant registered in the list of '100 of the world's worst invasive alien species'. An enrichment protocol was used to isolate microsatellite loci, and polymorphism was explored with 36 individuals collected in an invasive population from Australia and 20 individuals collected in a population from the native mainland location in South Africa. These primers showed a number of alleles per locus ranging from two to 10, and expected heterozygosities ranging from 0.083 to 0.826. Moreover, results of cross-species amplification are reported in five other Pheidole species and in seven other ants of the subfamily Myrmicinae.     

Behavioural and chemical mechanisms behind a Mediterranean ant–ant association

1. Interspecific competition among ants is common, and so is competitive exclusion among dominant ant species. In contrast, specific associations between non‐parasitic ant species are rare, especially in the temperate zones. As an exception, the subordinate ant Camponotus lateralis frequently co‐occurs with the dominant Crematogaster scutellaris but rarely with other dominant ants. 2. This association is one of various associations between Camponotus and Crematogaster species across the world. However, the mechanisms behind these co‐occurences are largely unknown. 3. In the present study, we therefore investigated the association of Ca. lateralis and Cr. scutellaris. We studied the spatial association of the nests, interspecific aggression, both species' cuticular hydrocarbon profiles, and their propensity to follow the other species' pheromone trails. 4. Crematogaster scutellaris usually attacked and displaced the generally submissive Ca. lateralis, but was significantly less aggressive at jointly used trails. Camponotus nests were always in close proximity to Crematogaster nests. 5. The cuticular hydrocarbons of both species consisted of alkanes with chain lengths between C21 and C35. The two species had 25 hydrocarbons in common, including mono‐, di‐, and tetramethyl alkanes. Despite this qualitative similarity, however, the quantitative hydrocarbon composition differed between the two species. 6. Camponotus lateralis followed artificial trails containing trail pheromones of Cr. scutellaris, but the latter did not follow Ca. lateralis trail pheromones. Interspecific trail‐following by Camponotus, but not vice versa, has been observed in another Camponotus–Crematogaster association and may be a more general mechanism that facilitates associations between the two ant genera.     

Intraspecific competition influences the symmetry and intensity of aggression in the Argentine ant

Cooperative social groups rely on the ability to distinguishmembers from nonmembers. Accordingly, social insects have evolveda variety of systems that allow discrimination of nest matesfrom non–nest mates. In this study, we show that experiencecan modify patterns of intraspecific aggression in Argentineants (Linepithema humile). In laboratory experiments, we foundthat aggression between colonies was often asymmetrical, butin all five cases, this asymmetry shifted to symmetrical aggressionafter contact with a hostile colony. Moreover, in the field,aggression between workers collected from colony borders wassymmetrical, whereas polarized aggression occurred between workerscollected 500 m away from colony borders. Coinciding with thisshift in aggression symmetry, we also observed an increase inboth the overall level of aggression and the frequency of aggressionin both the field and laboratory bioassays. We found littleevidence for colony-level competitive asymmetries stemming frompolarities in aggression at the worker level, either in thelaboratory or in the field. These results illustrate that recognitionsystems in Argentine ants are surprisingly dynamic and provideexperimental evidence for how recognition can be adjusted inresponse to specific circumstances—in this case the presenceof intraspecific competitors.     

Life-history stage and the population genetics of the tiger mosquito Aedes albopictus at a fine spatial scale

As a widespread vector of disease with an expanding range, the mosquito Aedes albopictus Skuse (Diptera: Culicidae) is a high priority for research and management. A. albopictus has a complex life history with aquatic egg, larval and pupal stages, and a terrestrial adult stage. This requires targeted management strategies for each life stage, coordinated across time and space. Population genetics can aid in A. albopictus control by evaluating patterns of genetic diversity and dispersal. However, how life stage impacts population genetic characteristics is unknown. We examined whether patterns of A. albopictus genetic diversity and differentiation changed with life stage at a spatial scale relevant to management efforts. We first conducted a literature review of field-caught A. albopictus population genetic papers and identified 101 peer-reviewed publications, none of which compared results between life stages. Our study uniquely examines population genomic patterns of egg and adult A. albopictus at five sites in Wake County, North Carolina, USA, using 8425 single nucleotide polymorphisms. We found that the level of genetic diversity and connectivity between sites varied between adults and eggs. This warrants further study and is critical for research aimed at informing local management.     

Isolation and characterization of polymorphic microsatellite loci for the invasive plant Cytisus scoparius

Scotch broom (Cytisus scoparius) is a serious weed of natural and pastoral ecosystems and is invasive in many countries worldwide. Eight polymorphic microsatellite loci were developed and characterized for this species. The number of alleles per locus ranged from five to 23. The range of observed heterozygosity was between 0.207 and 0.928. These microsatellite markers will be useful tools for studies of population genetics in the native and invasive range of this species.     

Detecting Nestmate Recognition Patterns in the Fission-Performing Ant <Emphasis Type="Italic">Aphaenogaster senilis</Emphasis>: A Comparison of Different Indices

In this paper, we compare three indices for nestmate recognition behaviors in the ant Aphaenogaster senilis within and among colonies and populations: the classical Aggression Index, a multidimensional Principal Component Analysis (PCA) Behavioral Index incorporating all observed behaviors, and a Chemical Profile Index for cuticular hydrocarbon composition. With these indices, we quantified nestmate recognition behaviors between workers from a parent nest and between those from independent colonies. The PCA Behavioral Index performed as well as or better than the classical Aggression Index in distinguishing small differences in nestmate recognition. Correlation analysis of the PCA Behavioral Index with the Chemical Profile Index may assist in identifying the occurrence of colony fissions. The PCA Behavioral Index correlated strongly with the Chemical Profile Index, suggesting that hydrocarbon profiling may be used as an alternative to nestmate recognition assays. This correlation also suggests that nestmate recognition in A. senilis is a graded response rather than a threshold one.     

Molecular systematics and population genetics of biological invasions: towards a better understanding of invasive species management

The study of population genetics of invasive species offers opportunities to investigate rapid evolutionary processes at work, and while the ecology of biological invasions has enjoyed extensive attention in the past, the recentness of molecular techniques makes their application in invasion ecology a fairly new approach. Despite this, molecular biology has already proved powerful in inferring aspects not only relevant to the evolutionary biologist but also to those concerned with invasive species management. Here, we review the different molecular markers routinely used in such studies and their application(s) in addressing different questions in invasion ecology. We then review the current literature on molecular genetic studies aimed at improving management and the understanding of invasive species by resolving of taxonomic issues, elucidating geographical sources of invaders, detecting hybridisation and introgression, tracking dispersal and spread and assessing the importance of genetic diversity in invasion success. Finally, we make some suggestions for future research efforts in molecular ecology of biological invasions.     

An invasive fish species within its native range: community effects and population dynamics of Gambusia affinis in the central United States

1. Management of invasive species benefits from detailed information on the biology of the invaders, both from where they have already invaded, and from within their areas of origin. Western mosquitofish, Gambusia affinis, is a widely invasive and destructive freshwater fish. However, within its native range, G. affinis co‐exists with many other fish species in a wide variety of habitats without obvious harm. 2. In this study, we used data on fish communities within the native range of G. affinis at 154 sites across a broad spatial scale to examine the effects of G. affinis on species richness and diversity of residual (species other than G. affinis) fish assemblages. We further used data based on annual samples at eight fixed river sites over 18 summers to examine temporal population dynamics of G. affinis and to test factors associated with population fluctuations. 3. Higher residual species richness occurred in the presence of G. affinis, but residual diversity did not differ. We found an inverse relationship between relative abundance of G. affinis and residual species richness (although effect size was extremely small), but no effect on residual diversity. 4. Gambusia affinis populations fluctuated markedly across summers at all eight fixed sites, but population sizes at a site over time were not autocorrelated. However, population fluctuations were highly correlated among sites across all years, suggesting that regional factors influenced population size. Regional abundance of G. affinis did not correlate with drought, rainfall or winter temperature, but varied with spring temperature. We suggest earlier onset of reproduction in warmer springs resulted in larger summer populations. 5. Overall, within its native range, G. affinis does not appear to impact negatively on the assemblages in which it occurs, possibly due to fluctuations in its density. These findings suggest that introduced Gambusia populations, and those of other invasive species, warrant careful monitoring over long periods of time where they have invaded. Long‐term monitoring of new populations can establish if they are prone to ‘boom and bust’ dynamics, in which case the invader may be less a threat than sometimes assumed. Population information from long‐term studies, either in their native ranges or at invaded sites, can thus help to form the basis of prudent, cost‐effective management strategies for invasive organisms.     

Inbreeding variability and population structure in the invasive haplodiploid palm‐seed borer (Coccotrypes dactyliperda)

We investigated the mating system and population genetic structure of the invasive haplodiploid palm‐seed borer Coccotrypes dactyliperda in California. We focused on whether these primarily inbreeding beetles have a ‘mixed‐breeding’ system that includes occasional outbreeding, and whether local inbreeding coefficients (F_IS) varied with dominant environmental factors. We also analysed the genetic structure of C. dactyliperda populations across local and regional scales. Based on the analysis of genetic variation at seven microsatellite loci in 1034 individual beetles from 59 populations, we found both high rates of inbreeding and plentiful evidence of mixed‐breeding. F_IS ranged from ?0.56 to 0.90, the highest variability reported within any animal species. There was a negative correlation between F_IS and latitude, suggesting that some latitude‐associated factor affecting mating decisions influenced inbreeding rates. Multiple regressions suggested that precipitation, but not temperature, may be an important correlate. Finally, we found highly significant genetic differentiation among sites, even over short geographic distances (< 1000 m).     

Molecular markers for systematic identification and population genetics of the invasive Ponto‐Caspian freshwater gammarid Dikerogammarus villosus (Crustacea,Amphipoda)

The Ponto‐Caspian amphipod, Dikerogammarus villosus, is an invasive species of many European rivers. First, we show that size difference of nrDNA ITS1 allows discriminating D. villosus from Dikerogammarus bispinosus, a closely related but morphologically hardly distinguishable species. Second, we present two types of polymorphic markers for D. villosus, three microsatellites and two single nucleotide polymorphisms (SNPs) of mtDNA COI gene, which were scored by polymerase chain reaction‐single strand conformational polymorphism (PCR‐SSCP). These markers will be very useful in studying population genetics of D. villosus.