Simon Dupont

Competition between invasive and indigenous species: an insular case study of subterranean termites

An important requirement for the management of invasive species is to identify the biological and ecological factors that influence the ability of such species to become established and spread within a new environment. Although competition is one of the key interactions determining the coexistence of species and exclusion, few studies directly examine the mechanism of competitive interactions within invasive communities. This study focused on putative competition in a social insect invader, R. flavipes, an American termite introduced into France, and an indigenous European termite, R. grassei. We first characterized and mapped a zone of sympatry between these two species. We then evaluated the degree of direct and indirect competition by comparing several life-history traits: behavioral aggression, chemical recognition and dispersion modes. Interspecific competition revealed that R. flavipes was dominant over R. grassei. Intraspecific competition was not found in R. flavipes while it appeared in varying degrees in R. grassei. These findings seemed to be correlated with the remarkable chemical homogeneity found in R. flavipes in comparison with R. grassei. Genetic analyses revealed that R. flavipes foraged over a greater distance than R. grassei colonies and might suggest a difference in the capacity to produce secondary reproductives. These findings suggest that R. flavipes has a significant advantage owing to competitive asymmetry that may enable the species to become dominant. The interspecific superiority, lack of intraspecific aggression and large extensive colonies, seem to be some of the reasons for its invasive success.

Global genetic analysis reveals the putative native source of the invasive termite, Reticulitermes flavipes, in France

Biological invasions are recognized as a major threat to both natural and managed ecosystems. Phylogeographic and population genetic analyses can provide information about the geographical origins and patterns of introduction and explain the causes and mechanisms by which introduced species have become successful invaders. Reticulitermes flavipes is a North American subterranean termite that has been introduced into several areas, including France where introduced populations have become invasive. To identify likely source populations in the USA and to compare the genetic diversity of both native and introduced populations, an extensive molecular genetic study was undertaken using the COII region of mtDNA and 15 microsatellite loci. Our results showed that native northern US populations appeared well differentiated from those of the southern part of the US range. Phylogenetic analysis of both mitochondrial and nuclear markers showed that French populations probably originated from southeastern US populations, and more specifically from Louisiana. All of the mtDNA haplotypes shared between the United States and France were found in Louisiana. Compared to native populations in Louisiana, French populations show lower genetic diversity at both mtDNA and microsatellite markers. These findings are discussed along with the invasion routes of R. flavipes as well as the possible mechanisms by which French populations have evolved after their introduction.

High occurrence of colony fusion in a European population of the American termite Reticulitermes flavipes

The coexistence of multiple unrelated reproductives within social insect colonies decreases the relatedness among colony members and therefore challenges kin selection theory. This study investigated the colony genetic structure of a French introduced population of the American subterranean termite Reticulitermes flavipes by analyzing genotypes at eight microsatellite loci and at one mtDNA region. Results revealed that all colonies contained numerous related secondary reproductives, and that 31% of colonies possessed more than two unrelated reproductives. The presence of several unrelated reproductives within colonies of this species is commonly assumed to result from colony fusion. Although such a high occurrence of colony fusion is the highest ever observed in a termite population, it is probable that the available methodology underestimated the detection of colony fusion in French populations. Overall, these results suggest that French colonies might differ strongly from the great majority of American colonies in their capacity to produce secondary reproductives as well as in their ability to merge. The nature and evolutionary origin of these population differences are discussed.

From speciation to introgressive hybridization: the phylogeographic structure of an island subspecies of termite, Reticulitermes lucifugus corsicus

BackgroundAlthough much research has been carried out into European Reticulitermes taxonomy in recent years, there is still much discussion about phylogenetic relationships. This study investigated the evolution from intra- to interspecific phylogeny in the island subspecies Reticulitermes lucifugus corsicus and threw new light on this phenomenon. An integrative approach based on microsatellites and mitochondrial and nuclear DNA sequences was used to analyze samples taken from a wide area around the Tyrrhenian sea and showed how the subspecies evolved from its origins to its most recent form on continental coasts.ResultsAccording to mitochondrial phylogeny and molecular clock calculations, island and continental taxa diverged significantly by vicariance in the Pleistocene glacial period. However, more recently, numerous migrations, certainly human-mediated, affected the structure of the populations. This study provided evidence of direct hybridization and multiple introgressions which occurred in several hybrid areas. Analysis using STRUCTURE based on microsatellite data identified a population in Provence (France) which differed considerably (Fst = 0.477) from populations on the island of Corsica and in Tuscany in the Italian peninsula. This new population, principally distributed in urban areas, is highly heterogeneous especially within the ITS2 regions where homogenization by concerted evolution does not appear to have been completed.ConclusionThis study provides an unusual picture of genetic interaction between termite populations in the Tyrrhenian area and suggests that more attention should be paid to the role of introgression and human impact on the recent evolution of European termites.

Geographical distribution, genetic diversity and social organization of a new European termite, Reticulitermes urbis (Isoptera: Rhinotermitidae)

Reticulitermes urbis is a recently described termite species that has probably been introduced into Western Europe where it has been found exclusively in urban areas. However, little is known about the geographic distribution and origin of R. urbis. This study was undertaken to determine whether this species was introduced from the Balkans. A parsimony network did not show any association between mtDNA haplotypes and geographic regions suggesting that western European populations were the result of human-mediated dispersion. Variation patterns of the COI and COII regions as well as microsatellites showed that the genetic diversity of Western European colonies was lower than for colonies collected in the Balkans, suggesting that the introduced populations suffered from a founder effect. As observed in the introduced populations of R. flavipes, all colonies of R. urbis had an extended-family structure with several reproductives. These results support the scenario that this termite was introduced into Western Europe.

Relationship between invasion success and colony breeding structure in a subterranean termite

Factors promoting the establishment and colonization success of introduced populations in new environments constitute an important issue in biological invasions. In this context, the respective role of pre‐adaptation and evolutionary changes during the invasion process is a key question that requires particular attention. This study compared the colony breeding structure (i.e. number and relatedness among reproductives within colonies) in native and introduced populations of the subterranean pest termite, Reticulitermes flavipes. We generated and analysed a data set of both microsatellite and mtDNA loci on termite samples collected in three introduced populations, one in France and two in Chile, and in the putative source population of French and Chilean infestations that has recently been identified in New Orleans, LA. We also provided a synthesis combining our results with those of previous studies to obtain a global picture of the variation in breeding structure in this species. Whereas most native US populations are mainly composed of colonies headed by monogamous pairs of primary reproductives, all introduced populations exhibit a particular colony breeding structure that is characterized by hundreds of inbreeding reproductives (neotenics) and by a propensity of colonies to fuse, a pattern shared uniquely with the population of New Orleans. These characteristics are comparable to those of many invasive ants and are discussed to play an important role during the invasion process. Our finding that the New Orleans population exhibits the same breeding structure as its related introduced populations suggests that this native population is pre‐adapted to invade new ranges.

Population Diversity in Cuticular Hydrocarbons and mtDNA in a Mountain Social Wasp

Nestmate recognition is a common phenomenon in social insects that typically is mediated by cuticular hydrocarbons. Geographical variation in cuticular hydrocarbons has been observed, although the pattern of variation is not consistent across species and is usually related to the biology and ecology of the different species. Polistes biglumis (Hymenoptera: Vespidae) is a social wasp that lives in high mountains where populations are separated by significant geographical barriers. Here we investigated the level of chemical variation among populations of P. biglumis in the Alps, and shed light on the phylogeography of this species. Populations could be discriminated by means of their cuticular hydrocarbon profiles, which showed a pattern consistent with the isolation-by-distance hypothesis. Molecular data highlighted two areas with different levels of haplotype diversity, although all wasps belonged to the same species. These results suggest that the populations of P. biglumis in the Alps are geographically isolated from one another, favoring their genetic and chemical differentiation.

Colony Breeding Structure of the Invasive Termite Reticulitermes urbis (Isoptera: Rhinotermitidae)

ABSTRACT Invasive species cause severe environmental and economic problems. The invasive success of social insects often appears to be related to their ability to adjust their social organization to new environments. To gain a better understanding of the biology of invasive termites, this study investigated the social organization of the subterranean termite, Reticulitermes urbis, analyzing the breeding structure and the number of reproductives within colonies from three introduced populations. By using eight microsatellite loci to determine the genetic structure, it was found that all the colonies from the three populations were headed by both primary reproductives (kings and queens) and secondary reproductives (neotenics) to form extended-family colonies. R. urbis appears to be the only Reticulitermes species with a social organization based solely on extended-families in both native and introduced populations, suggesting that there is no change in their social organization on introduction. F-statistics indicated that there were few neotenics within the colonies from urban areas, which did not agree with results from previous studies and field observations. This suggests that although several neotenics may be produced, only few become active reproductives. The results also imply that the invasive success of R. urbis may be based on different reproductive strategies in urban and semiurbanized areas. The factors influencing an individual to differentiate into a neotenic in Reticulitermes species are discussed.

Subterranean termite phylogeography reveals multiple postglacial colonization events in southwestern Europe

Abstract A long‐standing goal of evolutionary biology is to understand how paleoclimatic and geological events shape the geographical distribution and genetic structure within and among species. Using a diverse set of markers (cuticular hydrocarbons, mitochondrial and nuclear gene sequences, microsatellite loci), we studied Reticulitermes grassei and R. banyulensis, two closely related termite species in southwestern Europe. We sought to clarify the current genetic structure of populations that formed following postglacial dispersal from refugia in southern Spain and characterize the gene flow between the two lineages over the last several million years. Each marker type separately provided a fragmented picture of the evolutionary history at different timescales. Chemical analyses of cuticular hydrocarbons and phylogenetic analyses of mitochondrial and nuclear genes showed clear separation between the species, suggesting they diverged following vicariance events in the Late Miocene. However, the presence of intermediate chemical profiles and mtDNA introgression in some Spanish colonies suggests ongoing gene flow. The current genetic structure of Iberian populations is consistent with alternating isolation and dispersal events during Quaternary glacial periods. Analyses of population genetic structure revealed postglacial colonization routes from southern Spain to France, where populations underwent strong genetic bottlenecks after traversing the Pyrenees resulting in parapatric speciation.

A scientific note about a parasitoid that can parasitize the yellow-legged hornet, Vespa velutina nigrithorax, in Europe

The yellow-legged hornet, Vespa velutina nigrithorax, was accidentally introduced from China to southwestern France in 2004 (Haxaire et al. 2006). The species is currently colonizing France and neighboring countries (Spain, Portugal, Belgium, and Italy) (Darrouzet and Gevar 2012; Monceau et al. 2013). V. velutina is a pest in France because it preys upon domestic honeybees, Apis mellifera. Beekeeping operations are directly affected by this predation, with some beekeepers reporting colony losses (Darrouzet and Gevar 2012; Monceau et al. 2013).While many factors may play a role in the successful establishment of an invasive species, research has focused on the effects of parasites (Prenter et al. 2004). It has previously been suggested that invasive species are subject to less pressure from parasites than are sympatric native species because invasive species leave their parasites behind when they invade new habitats (enemy release hypothesis). They can thus allocate more resources to gr ...