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. K E Y W O R D S :Attractiveness, geographic isolation, Leptopilina clavipes, sperm storage, thelytoky, Wolbachia.

Whereas sexual reproduction may facilitate adaptation to complex environments with many biotic interactions, simplified environments are expected to favour asexual reproduction. In agreement with this, recent studies on invertebrates have shown a prevalence of asexual species in agricultural (simplified) but not in natural (complex) environments. We investigated whether the same correlation between reproductive mode and habitat can be found in different populations within one species. The parasitoid wasp Tetrastichus coeruleus forms an ideal model to test this question, since it occurs both in natural and agricultural environments. Further, we investigated whether Wolbachia infection caused parthenogenesis in female-biased populations. In contrast to the general pattern, in Dutch and French natural areas, we found Wolbachia-infected, highly female-biased populations that reproduce parthenogenetically. In contrast, populations on Dutch agricultural fields were not infected with Wolbachia, showed higher frequencies of males and reproduced sexually. However, we also found a female-only, Wolbachia-infected population on agricultural fields in north-eastern United States. All Wolbachia-infected populations were infected with the same Wolbachia strain. At this moment, we do not have a convincing explanation for this deviation from the general pattern of ecology and reproductive mode. It may be that asparagus agricultural fields differ from other crop fields in ways that favour sexual reproduction. Alternatively, Wolbachia may manipulate life history traits in its host, resulting in different fitness pay-offs in different habitats. The fixation of Wolbachia in the United States populations (where the species was introduced) may be due to founder effect and lack of uninfected, sexual source populations.

Wolbachia are endosymbiotic bacteria known to manipulate the reproduction of their hosts. Some populations of the parasitoid wasp Asobara japonica are infected with Wolbachia and reproduce parthenogenetically, while other populations are not infected and reproduce sexually. Wolbachia-infected A. japonica females regularly produce small numbers of male offspring. Because all females in the field are infected and infected females are not capable of sexual reproduction, male production seems to be maladaptive. We investigated why these females nevertheless produce males. We tested three hypotheses: high rearing temperatures could result in higher offspring sex ratios (more males), low Wolbachia titer of the mother could lead to higher offspring sex ratios and/or the Wolbachia infection is of relatively recent origin and not enough time has passed to allow complete coadaptation between Wolbachia and host. In all, 33% of the Wolbachia-infected females produced males and 56% of these males were also infected with Wolbachia. Neither offspring sex ratio nor male infection frequency was significantly affected by rearing temperature or Wolbachia concentration of the mother. The mitochondrial DNA sequence of one of the uninfected populations was identical to that of two of the infected populations. Therefore, the initial Wolbachia infection of A. japonica must have occurred recently. Mitochondrial sequence variation among the infected populations suggests that the spread of Wolbachia through the host populations involved horizontal transmission. We conclude that the occasional male production by Wolbachia-infected females is most likely a maladaptive side effect of incomplete coevolution between symbiont and host in this relatively young infection.

Wolbachia is a maternally inherited bacterium that manipulates the reproduction of its host. Recent studies have shown that male-killing strains can induce cytoplasmic incompatibility (CI) when introgressed into a resistant host. Phylogenetic studies suggest that transitions between CI and other Wolbachia phenotypes have also occurred frequently, raising the possibility that latent CI may be widespread among Wolbachia. Here, we investigate whether a parthenogenesis-inducing Wolbachia strain can also induce CI. Parthenogenetic females of the parasitoid wasp Asobara japonica regularly produce a small number of males that may be either infected or not. Uninfected males were further obtained through removal of the Wolbachia using antibiotics and from a naturally uninfected strain. Uninfected females that had mated with infected males produced a slightly, but significantly more male-biased sex ratio than uninfected females that had mated with uninfected males. This effect was strongest in females that mated with males that had a relatively high Wolbachia titer. Quantitative PCR indicated that infected males did not show higher ratios of nuclear versus mitochondrial DNA content. Wolbachia therefore does not cause diploidization of cells in infected males. While these results are consistent with CI, other alternatives such as production of abnormal sperm by infected males cannot be completely ruled out. Overall, the effect was very small (9%), suggesting that if CI is involved it may have degenerated through the accumulation of mutations.

Wolbachia are endosymbiotic bacteria known to manipulate the reproduction of their hosts. These manipulations are expected to have consequences on the population genetics of the host, such as heterozygosity levels, genetic diversity and gene flow. The parasitoid wasp Tetrastichus coeruleus has populations that are infected with parthenogenesis-inducing Wolbachia and populations that are not infected. We studied the population genetics of T. coeruleus between and within Wolbachia-infected and uninfected populations, using nuclear microsatellites and mitochondrial DNA. We expected reduced genetic diversity in both DNA types in infected populations. However, migration and gene flow could introduce new DNA variants into populations. We therefore paid special attention to individuals with unexpected (genetic) characteristics. Based on nuclear and mitochondrial DNA, two genetic clusters were evident: a thelytokous cluster containing all Wolbachia-infected, parthenogenetic populations and an arrhenotokous cluster containing all uninfected, sexual populations. Nuclear and mitochondrial DNA did not exhibit concordant patterns of variation, although there was reduced genetic diversity in infected populations for both DNA types. Within the thelytokous cluster, there was nuclear DNA variation, but no mitochondrial DNA variation. This nuclear DNA variation may be explained by occasional sex between infected females and males, by horizontal transmission of Wolbachia, and/or by novel mutations. Several females from thelytokous populations were uninfected and/or heterozygous for microsatellite loci. These unexpected characteristics may be explained by migration, by inefficient transmission of Wolbachia, by horizontal transmission of Wolbachia, and/or by novel mutations. However, migration has not prevented the build-up of considerable genetic differentiation between thelytokous and arrhenotokous populations.

No abstract

The cricetid species Democricetodon gracilis (Fahlbusch, 1964), Democricetodon mutilus (Fahlbusch, 1964), Megacricetodon bavaricus (Fahlbusch, 1964), and Megacricetodon minor (Lartet, 1851) from Sandelzhausen (Early/Middle Miocene boundary, MN5, Southern Germany) are represented by more than 2,000 molars providing substantial data on the variability in sizes and on variation in morphologies. Temporal and spatial distributions of these species indicate that they are probably immigrants in SE Germany, originating from south-eastern or more southern areas. Except for M. minor, whose origin is not clear and could be regarded as an immigrant from more eastern areas. Using the length of the mesoloph in the second upper molars as an indicator of habitat, D. gracilis and M. minor had a preference for a humid habitat, D. mutilus and M. bavaricus a preference for drier habitats.Kurzfassung Die Hamster-Arten Democricetodon gracilis (Fahlbusch, 1964), Democricetodon mutilus (Fahlbusch, 1964), Megacricetodon bavaricus (Fahlbusch, 1964) und Megacricetodon minor (Lartet, 1851) sind durch mehr als 2000 Zähne in Sandelzhausen (Grenzbereich Unter-/Mittelmiozän, MN5, Süddeutschland) belegt und liefern umfangreiche Daten zur Kenntnis der Variabilität der Größe sowie der Morphologie. Die zeitliche und räumliche Verbreitung dieser fossilen Arten zeigen, dass sie wahrscheinlich aus südöstlichen oder mehr südlichen Gebieten nach Deutschland eingewandert sind. Eine Ausnahme könnte M. minor darstellen, dessen Herkunft nicht klar ist und vielleicht als ein Einwanderer aus östlichen Gebieten betrachtet werden könnte. Die Länge des Mesoloph am zweiten oberen Molar erlaubt Rückschlüsse auf bevorzugte Lebensräume und ergibt für D. gracilis und M. minor eine Vorliebe für ein eher feuchtes Habitat während D. mutilus und M. bavaricus offensichtlich trockenere Habitate bevorzugten. Schlü sselwörter Cricetidae Á Democricetodon Á Megacricetodon Á Deutschland Á MN5 Á Verbreitung Á Palökologie

Wolbachia are endosymbiotic bacteria known to manipulate the reproduction of their hosts by, for example, inducing parthenogenesis. In most cases of Wolbachia-induced parthenogenesis, the infection is fixed and the entire host population consists of females. In the absence of males and sexual reproduction, genes involved in sexual reproduction are not actively maintained by selection. Accumulation of neutral mutations or selection against maintenance of sexual traits may lead to their loss or deterioration. In addition, females may lose the ability to reproduce sexually due to 'functional virginity mutations' that may spread concomitantly with the Wolbachia infection through a population. The parasitoid wasp Tetrastichus coeruleus (Nees) (Hymenoptera: Eulophidae) forms an ideal model to study the decay of sexual functionality, because it has both Wolbachia-infected, parthenogenetic populations and uninfected, sexual populations. We compared several components of sexual functionality of arrhenotokous (sexual) and thelytokous (parthenogenetic) T. coeruleus females. First, we tested whether arrhenotokous and thelytokous females were equally attractive and receptive to males. Second, we examined whether mating is costly to females by measuring the life span of mated and virgin females. Last, we studied the morphology of the spermathecae of arrhenotokous and thelytokous females. Mated females had shorter life spans than virgin females, showing that mating carried a fitness cost. Two sexual traits of thelytokous females have degraded compared to arrhenotokous females. Arrhenotokous and thelytokous females were equally attractive to males, but thelytokous females were unreceptive to males. Furthermore, there was a clear difference in spermathecal morphology between arrhenotokous and thelytokous females. Our data do not allow distinction between the various potential causes of such degradation. Although the longevity cost of mating may indicate selection against the maintenance of costly sexual traits, accumulation of neutral mutations, functional virginity mutations, manipulation by Wolbachia, and/or the genetic distance between the two populations may all have contributed to the decay of sexual traits in thelytokous females.