In Europe the most abundant naturally residing termite is the subterranean genus Reticulitermes (Rhinotermitidae). Six phenotypes of Reticulitermes have been identified on the basis of morphological, chemical (cuticular hydrocarbons and soldier defensive secretions), and molecular (enzymatic alleles and mitochondrial ND1 sequence) features. They are R. santonensis in western France, R. grassei in southwestern France, northwestern and southern Spain and Portugal, R. banyulensis in northeastern Spain, central area of the Iberian Peninsula and southwestern France, R. lucifugus in Italy and southeastern France, R. balkanensis in the Balkans and R. sp. nov., a recently identified urban phenotype resembling R. balkanensis, in northern Italy and southeastern France. R. santonensis is close kin to the American species R. flavipes. R. grassei, R. banyulensis and R. lucifugus belong to the same species complex. R. balkanensis and the new phenotype R. sp. nov. are close to R. santonensis regarding cuticular hydrocarbons, to the lucifugus complex regarding DNA and to R. clypeatus from Israel regarding morphology. The species status of these genotypes has been confirmed by the mechanisms of species isolation. Prevention of hybridization depends on the method of colony formation in each species. Swarming dates, differences in pheromones, and infertility prevent hybridization by sexual alates. Interspecific aggression between workers prevents hybridization by neotenics. Behavioral and molecular studies have provided many data on the genetic structure of nests, which varies according to species and location. All colonies of R. santonensis are open all year. The colonies of R. grassei in southern areas and all colonies of R. banyulensis are closed families with generally a single reproductive couple. The colonies of R. grassei in northern areas and the colonies of R. lucifugus are open in the summer and closed in the winter. Based on the here presented data, the taxonomy and the speciation of the Reticulitermes genus in Europe are discussed.

Reticulitermes santonensis is a subterranean termite that invades urban areas in France and elsewhere where it causes damage to human-built structures. We investigated the breeding system, colony and population genetic structure, and mode of dispersal of two French populations of R. santonensis. Termite workers were sampled from 43 and 31 collection points, respectively, from a natural population in west-central France (in and around the island of Oleron) and an urban population (Paris). Ten to 20 workers per collection point were genotyped at nine variable microsatellite loci to determine colony identity and to infer colony breeding structure. There was a total of 26 colonies, some of which were spatially expansive, extending up to 320 linear metres. Altogether, the analysis of genotype distribution, F-statistics and relatedness coefficients suggested that all colonies were extended families headed by numerous neotenics (nonwinged precocious reproductives) probably descended from pairs of primary (winged) reproductives. Isolation by distance among collection points within two large colonies from both populations suggested spatially separated reproductive centres with restricted movement of workers and neotenics. There was a moderate level of genetic differentiation (F(ST) = 0.10) between the Oleron and Paris populations, and the number of alleles was significantly higher in Oleron than in Paris, as expected if the Paris population went through bottlenecks when it was introduced from western France. We hypothesize that the diverse and flexible breeding systems found in subterranean termites pre-adapt them to invade new or marginal habitats. Considering that R. santonensis may be an introduced population of the North American species R. flavipes, a breeding system consisting primarily of extended family colonies containing many neotenic reproductives may facilitate human-mediated spread and establishment of R. santonensis in urban areas with harsh climates.

Nestmate recognition plays a key role in kin selection to maintain colony integrity in social insects. Previous studies have demonstrated that nestmate recognition is dependent on detection of cuticular hydrocarbons. However, the absence of intraspecific aggression between some colonies of Isoptera and social Hymenoptera questions whether kin recognition must occur in social insects. The purpose of this study was to determine if cuticular hydrocarbon similarity and high genetic relatedness could explain the lack of intraspecific aggression among and within colonies of the introduced subterranean termite Reticulitermes santonensis. We performed both GC analysis of cuticular hydrocarbons and genotyping by using 10 DNA microsatellite loci on the same 10 workers from each of 14 parisian colonies. Multivariate analyses demonstrated correspondence between cuticular hydrocarbon patterns and genetic variation. By using a redundancy analysis combining chemical and genetic data, we found that a few hydrocarbons (mainly short vs. long chains; saturated vs. unsaturated alkanes) were associated with most genetic variation. We also found a strong positive correlation between chemical and genetic distances between colonies, thus providing evidence of a genetic basis for cuticular hydrocarbon variation. However, genetic distance did not account for all chemical variation, thus suggesting that some hydrocarbon variation was environmentally derived. Investigation at the intracolony level indicated that cuticular hydrocarbons did not depend on colony social structure. Based on our findings, we speculate that the absence of intraspecific aggression in R. santonensis may result from a loss of diversity in genetically derived recognition compounds in this species that presumably descended from R. flavipes populations imported from North America.