Clonal genetic variation in a Wolbachia-infected asexual wasp: horizontal transmission or historical sex?

Wolbachia are endocellular bacteria known for manipulating the reproductive systems of many of their invertebrate hosts. Wolbachia are transmitted vertically from mother to offspring. In addition, new infections result from horizontal transmission between different host species. However, to what extent horizontal transmission plays a role in the spread of a new infection through the host population is unknown. Here, we investigate whether horizontal transmission of Wolbachia can explain clonal genetic variation in natural populations of Leptopilina clavipes, a parasitoid wasp infected with a parthenogenesis-inducing Wolbachia. We assessed variance of markers on the nuclear, mitochondrial and Wolbachia genomes. The nuclear and mitochondrial markers displayed significant and congruent variation among thelytokous wasp lineages, showing that multiple lineages have become infected with Wolbachia. The alternative hypothesis in which a single female became infected, the daughters of which mated with males (thus introducing nuclear genetic variance) cannot account for the presence of concordant variance in mtDNA. All Wolbachia markers, including the hypervariable wsp gene, were invariant, suggesting that only a single strain of Wolbachia is involved. These results show that Wolbachia has transferred horizontally to infect multiple female lineages during the early spread through L. clavipes. Remarkably, multiple thelytokous lineages have persisted side by side in the field for tens of thousands of generations.

Effects of parthenogenesis and geographic isolation on female sexual traits in a parasitoid wasp.

Population divergence in sexual traits is affected by different selection pressures, depending on the mode of reproduction. In allopatric sexual populations, aspects of sexual behavior may diverge due to sexual selection. In parthenogenetic populations, loss-of-function mutations in genes involved in sexual functionality may be selectively neutral or favored by selection. We assess to what extent these processes have contributed to divergence in female sexual traits in the parasitoid wasp Leptopilina clavipes in which some populations are infected with parthenogenesis-inducing Wolbachia bacteria. We find evidence consistent with both hypotheses. Both arrhenotokous males and males derived from thelytokous strains preferred to court females from their own population. This suggests that these populations had already evolved population-specific mating preferences when the latter became parthenogenetic. Thelytokous females did not store sperm efficiently and fertilized very few of their eggs. The nonfertility of thelytokous females was due to mutations in the wasp genome, which must be an effect of mutation accumulation under thelytoky. Divergence in female sexual traits of these two allopatric populations has thus been molded by different forces: independent male/female coevolution while both populations were still sexual, followed by female-only evolution after one population switched to parthenogenesis.

Spatial components of bird diversity in the Andes of Colombia: implications for designing a regional reserve system.

Beta diversity, or the turnover in species composition among sampling sites in a region, is an important criterion for obtaining adequate representation of regional biodiversity in systems of protected areas. Recently, the additive model for partitioning regional (gamma) diversity (in opposition to the multiplicative model) has been proposed because it allows a direct measure of the contribution of beta diversity to gamma diversity. We determined avian beta diversity along latitudinal (among neighboring river drainages) and elevational axes in a 1347-km2 region on the western slope of the Central Cordillera of the Colombian Andes, where a regional system of protected areas is being designed. We then compared avian beta diversity between sites based on rapid versus long-term (>1 year) inventories and between fragmented sites versus continuous forest. Overall, beta diversity represented 63.1% of gamma diversity among 16 sites. Elevational differences in species composition accounted for 43.3% of regional diversity, whereas differences among drainages accounted for 19.8%. A complementary cluster analysis showed that sites grouped by elevational zones. Rapid inventories overestimated beta diversity because of sampling effects, but the effect was biologically small. Estimators of species richness derived from species accumulation curves provided a useful alternative to compensate for undersampling in short-term surveys. Forest fragmentation increased beta diversity because of differential local extinction of populations. Nevertheless, in our region, forest fragments contributed to gamma diversity because they contained complementary sets of species. More importantly, they contained populations of special-interest species. Although the region is relatively small, our analyses indicate that spatial differentiation of the biota is an important factor for deciding number and location of protected areas in the Andean region.