COMPARATIVE GEOGRAPHIC STRUCTURES OF TWO PARASITOID-HOST INTERACTIONS

Recent studies of parasitoid-host interactions have demonstrated that parasitoids and their hosts are geographically structured for traits such as virulence and encapsulation defenses, but no studies have yet compared the geographic structure of parasitoids and hosts using neutral genetic markers. Such studies of geographic structure are needed to evaluate the underlying geographic scale at which these interactions evolve and allow assessment of the relative effects of selection and gene flow on the geographic structure observed in traits under selection. We used sequence data from the mitochondrial DNA cytochrome oxidase I and II subunits to document and compare the geographic structures of the parasitoid Agathis thompsoni and its moth host Greya subalba. We also documented the geographic structure of G. enchrysa and compared it to the geographic structure of its parasitoid Agathis n. sp. The results demonstrated that parasitoids and their hosts may have incongruent patterns of geographic structure as assessed by molecular markers. As a consequence, the geographic scale at which the interaction evolves may be different for each species involved in the interaction. Depending on the interplay of selection and gene flow, there may not be a one-to-one correspondence of traits important in the interaction between parasitoids and their hosts at the level of local populations. The geographic structures of A. thompsoni and G. subalba and Agathis n. sp. and G. enchrysa provide further evidence of the potential importance of the formation of geographic mosaics in coevolving parasitoid-host interactions and evolving interactions in general.     

One step ahead: a parasitoid disperses farther and forms a wider geographic population than its fig wasp host

The structure of populations across landscapes influences the dynamics of their interactions with other species. Understanding the geographic structure of populations can thus shed light on the potential for interacting species to co‐evolve. Host–parasitoid interactions are widespread in nature and also represent a significant force in the evolution of plant–insect interactions. However, there have been few comparisons of population structure between an insect host and its parasitoid. We used microsatellite markers to analyse the population genetic structure of Pleistodontes imperialis sp. 1, a fig‐pollinating wasp of Port Jackson fig (Ficus rubiginosa), and its main parasitoid, Sycoscapter sp. A, in eastern Australia. Besides exploring this host–parasitoid system, our study also constitutes, to our knowledge, the first study of population structure in a nonpollinating fig wasp species. We collected matched samples of pollinators and parasitoids at several sites in two regions separated by up to 2000 km. We found that pollinators occupying the two regions represent distinct populations, but, in contrast, parasitoids formed a single population across the wide geographic range sampled. We observed genetic isolation by distance for each species, but found consistently lower F_ST and R_ST values between sites for parasitoids compared with pollinators. Previous studies have indicated that pollinators of monoecious figs can disperse over very long distances, and we provide the first genetic evidence that their parasitoids may disperse as far, if not farther. The contrasting geographic population structures of host and parasitoid highlight the potential for geographic mosaics in this important symbiotic system.     

Coevolution across landscapes: a spatially explicit model of parasitoid-host coevolution

The geographic mosaic theory of coevolution suggests that population spatial structure may have a strong impact on coevolutionary dynamics. Therefore, coevolution must be studied across geographic scales, not just in single populations. To examine the impact of movement rate on coevolutionary dynamics, we developed a spatially explicit model of host–parasitoid coevolution. We described space as a coupled-map lattice and assumed that resistance (defined as the ability of a host to encapsulate a parasitoid egg) and virulence (defined as the successful parasitization of a host) traits were graded and costly. The model explicitly detailed population and evolutionary dynamics. When holding all parameters constant and varying only the movement rate of the host and parasitoid, profoundly different dynamics were observed. We found that fluctuations in the mean levels of resistance and virulence in the global population were greatest when the movement rate of the host and parasitoid was high. In addition, we found that the variation in resistance and virulence levels among neighboring patches was greatest when the movement rates of the host and parasitoid was low. However, as the distance among patches increased, so did the variation in resistance and virulence levels regardless of movement rate. These generalizations did not hold when spatial patterns in the distribution of resistance and virulence traits, such as spirals, were observed. Finally, we found that the evolution of resistance and virulence caused the abundance of hosts to increase and the abundance of parasitoids to decrease. As a result, the spatial distribution of hosts and parasitoids was influenced.     

Partitioning of herbivore hosts across time and food plants promotes diversification in the Megastigmus dorsalis oak gall parasitoid complex

Communities of insect herbivores and their natural enemies are rich and ecologically crucial components of terrestrial biodiversity. Understanding the processes that promote their origin and maintenance is thus of considerable interest. One major proposed mechanism is ecological speciation through host‐associated differentiation (HAD), the divergence of a polyphagous species first into ecological host races and eventually into more specialized daughter species. The rich chalcid parasitoid communities attacking cynipid oak gall wasp hosts are structured by multiple host traits, including food plant taxon, host gall phenology, and gall structure. Here, we ask whether the same traits structure genetic diversity within supposedly generalist parasitoid morphospecies. We use mitochondrial DNA sequences and microsatellite genotypes to quantify HAD for Megastigmus (Bootanomyia) dorsalis, a complex of two apparently generalist cryptic parasitoid species attacking oak galls. Ancient Balkan refugial populations showed phenological separation between the cryptic species, one primarily attacking spring galls, and the other mainly attacking autumn galls. The spring species also contained host races specializing on galls developing on different host‐plant lineages (sections Cerris vs. Quercus) within the oak genus Quercus. These results indicate more significant host‐associated structuring within oak gall parasitoid communities than previously thought and support ecological theory predicting the evolution of specialist lineages within generalist parasitoids. In contrast, UK populations of the autumn cryptic species associated with both native and recently invading oak gall wasps showed no evidence of population differentiation, implying rapid recruitment of native parasitoid populations onto invading hosts, and hence potential for natural biological control. This is of significance given recent rapid range expansion of the economically damaging chestnut gall wasp, Dryocosmus kuriphilus, in Europe.     

Comparative population genetic structures of the fruit fly Urophora cardui and its primary parasitoid Eurytoma robusta

The interaction between two species may depend on geographic scale and this in turn can affect co‐evolution among them. The present study comparatively examines population genetic structures of the tephritid gall fly Urophora cardui and its primary ectoparasitoid Eurytoma robusta for inference of relative dispersal patterns and host‐parasitoid specificity. Genetic differentiation patterns indicated two levels of hierarchical structure in both species: locally similar distance‐dependencies but globally differences. Locally, both species showed isolation by distance and a high correlation between host and parasitoid F_ST for the same population‐pairs was found. At the local level, E. robusta populations were most structured. These findings suggest that locally E. robusta is tracking behind its host, U. cardui, and that colonisation of new patches by both species underlie a stepping‐stone dispersal process. The investigation as a whole showed that U. cardui populations were hierarchically structured across a genetic‐geographical cline. There was no sign of a comparable cline in E. robusta where populations globally became independent of one another and of the host. The different degree of hierarchical genetic structure of the two species suggests that dispersal processes or interactions differ relative to geographical scale and population history.     

Host–parasitoid community model and its potential application in biological control of cereal stemborers in Kenya

A two-host–two-parasitoid model was constructed to assess the effects of the introduced larval parasitoid, the braconid Cotesia flavipes, on its primary target host, the invasive crambid Chilo partellus, and on secondary host species, in inter-specific competition with Cotesia sesamiae, the main native parasitoid species of stemborers in Kenya. The model assumed that: (1) there was no host discrimination by either parasitoid species; (2) Cotesia flavipes was the superior competitor that out-competed Cotesia sesamiae when the host was suitable; and (3) Cotesia flavipes could only develop in an unsuitable host if it had been previously parasitized by Cotesia sesamiae. Model parameters were estimated from surveys conducted in Kenya and from laboratory experiments. Different scenarios of host and parasitoid species composition and host suitability occurring in the different ecological zones in Kenya were analyzed. Results indicated that: (1) the coexistence of stemborer host populations are determined by their population growth rates, the degree of aggregation of the parasitoids and their searching efficiency; (2) in the regions where both the invasive and the predominant native host species were suitable to either parasitoid species, stemborer densities would be reduced to and controlled at low densities, and Cotesia flavipes would become the dominant parasitoid species. However, the extinction or predominance of the native stemborer species depends on the ratio of the growth rates of exotic and native stemborers and their relative searching efficiencies; and (3) if the native host species was acceptable but unsuitable to Cotesia flavipes, the parasite would not become established.     

Molecular biogeography and host relations of a parasitoid fly

Successful geographic range expansion by parasites and parasitoids may also require host range expansion. Thus, the evolutionary advantages of host specialization may trade off against the ability to exploit new host species encountered in new geographic regions. Here, we use molecular techniques and confirmed host records to examine biogeography, population divergence, and host flexibility of the parasitoid fly, Ormia ochracea (Bigot). Gravid females of this fly find their cricket hosts acoustically by eavesdropping on male cricket calling songs; these songs vary greatly among the known host species of crickets. Using both nuclear and mitochondrial genetic markers, we (a) describe the geographical distribution and subdivision of genetic variation in O. ochracea from across the continental United States, the Mexican states of Sonora and Oaxaca, and populations introduced to Hawaii; (b) demonstrate that the distribution of genetic variation among fly populations is consistent with a single widespread species with regional host specialization, rather than locally differentiated cryptic species; (c) identify the more‐probable source populations for the flies introduced to the Hawaiian islands; (d) examine genetic variation and substructure within Hawaii; (e) show that among‐population geographic, genetic, and host song distances are all correlated; and (f) discuss specialization and lability in host‐finding behavior in light of the diversity of cricket songs serving as host cues in different geographically separate populations.     

Behavioural differences between Aphidius ervi populations from two tritrophic systems are due to phenotypic plasticity

The Palaeoarctic parasitoid Aphidius ervi Haliday (Hymenoptera, Aphidiidae) parasitises legume aphids in its region of origin. In Chile, it parasitises both legume and cereal aphids. This special situation was studied at two levels: (i) the host searching behaviour of A. ervi from two different tritrophic systems (Acyrthosiphon pisum on alfalfa and Sitobion avenae on wheat) was investigated in dual choice tests in a wind tunnel between odours from both A. pisum-alfalfa host plant complex (HPC) and S. avenae-wheat HPC, and (ii) the genetic structure of A. ervi populations from both sources using molecular markers. Responses of A. ervi females to volatile olfactory cues emanating from A. pisum-alfalfa HPC and S. avenae-wheat HPC were significantly higher towards the HPC on which they were reared during the last generation before experimentation, regardless of the origin of the parasitoid. As previously described for this parasitoid species, oviposition experience was also of major relevance in the preferences of female parasitoids. On the other hand, variation in mitochondrial DNA segments and RAPD-PCR polymorphism using total DNA showed the absence of host-based population structure and a high genetic homogeneity between these A. ervi populations. These results reject the possible existence of different host-strains of this parasitoid in Chile.     

Comparative phylogeography across two trophic levels: the oak gall wasp Andricus kollari and its chalcid parasitoid Megastigmus stigmatizans

Insect parasitoids are important components of many terrestrial ecosystems. However, relatively little is known about the mechanisms responsible for structuring their populations. Here we investigate the ability of Megastigmus stigmatizans, an oak gall wasp parasitoid, to track its host Andricus kollari over two different timescales, and examine its current population structure across a divide in host population structure. The divide represents a transition in gall wasp host-plant species and offers the opportunity to examine whether the split, which divides gall wasp populations, manifests itself in the next trophic level. Analysis of mitochondrial haplotype data for parasitoid and host reveals: (i) A similar phylogeographic population structure for both, with Iberian populations more derived with respect to more eastern populations. (ii) It is likely that the host colonized the Iberian refuge earlier than the parasitoid, probably by at least one glacial cycle. (iii) Recent range expansion of central European host populations northwards has resulted in pursuit by parasitoids from the same geographic origin. (iv) In addition, Iberian parasitoid populations have crossed a major divide in host population structure to invade northern Europe. Such human-facilitated escape from natural refugial distributions may have important implications for the composition and structure of northern European gall wasp communities.     

Host selection behaviour and specificity of the solitary parasitoid of mealybugs Anagyrus sp. nr. pseudococci (Girault) (Hymenoptera,Encyrtidae)

The host selection behaviour of Anagyrus sp. nr. pseudococci was compared in no-choice tests among five mealybug species of different geographical and phylogenetic origin, including the Mediterranean native host, Planococcus ficus and four exotic mealybug species, one of the same genus, Pl. citri, two Pseudococcus species, Ps. calceolariae and Ps. viburni and a more distant one, Phenacoccus peruvianus. All five studied mealybug species were recognised by the parasitoid as potential hosts and parasitised, but the behavioural pattern of host recognition, host handling and the level of host acceptance of Anagyrus sp. nr. pseudococci significantly varied among the five studied species, indicating a clear preference for the two Planococcus species, Pl. ficus in particular. The results suggest that A. sp. nr. pseudococci has a broader host range and a more generalist behaviour in comparison with other Anagyrus species. Practical implications of the findings are discussed.     

Selection of Bemisia nymphal stages for oviposition or feeding,and host-handling times of arrhenotokous and thelytokous <Emphasis Type="Italic">Eretmocerus mundus</Emphasis> and arrhenotokous <Emphasis Type="Italic">E. eremicus</Emphasis>

Host-handling behavior is an important aspect of parasitoid foraging behavior. When a parasitoid encounters a potential host, the handling behavior starts with the evaluation of the host and continues if the host has been judged acceptable. Host handling is usually terminated after egg laying or host feeding and host marking. Host-handling behavior of an arrhenotokous population of two Eretmocerus species, E. mundus Mercet and E. eremicus Rose and Zolnerowich, along with a thelytokous population of E. mundus were compared under laboratory conditions. Several elements of host-handling behavior, including encountering, ascending, turning on host, descending, preening, egg laying, and host feeding were recorded. There were no correlations among the durations of these phases across parasitoid populations/species or host nymphal instars. Duration of different phases of host-handling behavior showed only slight and sometimes significant differences between different Eretmoceruspopulations/species. The actual laying of the egg had the longest duration of all host-handling behaviors, and was longer on third nymphal instars than on younger ones. Females of the three populations/species accepted the first three nymphal stages either for egg laying or for host feeding. Females spent a lot of time to make wounds in the host when preparing for host feeding, and eventually killed the host. The implications of these findings for the use of the different Eretmoceruspopulations/species in biological control are discussed.     

The effect of multi‐species host density on superparasitism and sex ratio in a gregarious parasitoid

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Patterns of host utilisation by herbivore assemblages of the genus Caloptilia (Lepidoptera; Gracillariidae) on congeneric maple tree (Acer) species

1. The pattern of host utilisation by congeneric Caloptilia caterpillars on 14 different species of Acer (maple) was investigated in temperate mixed forests of central Japan. A multi‐filtering model of host plant utilisation was proposed to address how phylogenetically related herbivore assemblages are constructed on phylogenetically related host plant species. 2. Two hypotheses were examined. The first questioned whether a negative relationship exists between the phylogenetic distance of plants from the most suitable host species and the abundance of herbivorous insects on the host. Regarding the second, it was investigated whether the assemblage dissimilarity of herbivorous insects among host plant species increases with increasing distance of plant phylogeny and traits. 3. Mantel and partial Mantel tests were used to measure the relationship between assemblage dissimilarity of Caloptilia species and the distance of plant phylogeny and leaf traits. 4. Both hypotheses were confirmed, clearly suggesting that the utilisation and suitability of hosts for Caloptilia caterpillars were strongly influenced by phylogenetic relatedness and leaf trait similarity among Acer species. This implies that phylogenetic distance is an integrated measure of phenotypic and ecological attributes of congeneric Acer species that can be used to explain specialisation and constraints of host utilisation of congeneric herbivore species even on a short evolutionary timescale.     

Genome‐wide differentiation in closely related populations: the roles of selection and geographic isolation

Population divergence in geographic isolation is due to a combination of factors. Natural and sexual selection may be important in shaping patterns of population differentiation, a pattern referred to as ‘isolation by adaptation’ (IBA). IBA can be complementary to the well‐known pattern of ‘isolation by distance’ (IBD), in which the divergence of closely related populations (via any evolutionary process) is associated with geographic isolation. The barn swallow Hirundo rustica complex comprises six closely related subspecies, where divergent sexual selection is associated with phenotypic differentiation among allopatric populations. To investigate the relative contributions of selection and geographic distance to genome‐wide differentiation, we compared genotypic and phenotypic variation from 350 barn swallows sampled across eight populations (28 pairwise comparisons) from four different subspecies. We report a draft whole‐genome sequence for H. rustica, to which we aligned a set of 9493 single nucleotide polymorphisms (SNPs). Using statistical approaches to control for spatial autocorrelation of phenotypic variables and geographic distance, we find that divergence in traits related to migratory behaviour and sexual signalling, as well as geographic distance, together explain over 70% of genome‐wide divergence among populations. Controlling for IBD, we find 42% of genomewide divergence is attributable to IBA through pairwise differences in traits related to migratory behaviour and sexual signalling alone. By (i) combining these results with prior studies of how selection shapes morphological differentiation and (ii) accounting for spatial autocorrelation, we infer that morphological adaptation plays a large role in shaping population‐level differentiation in this group of closely related populations.     

DEFENSE TRAITS OF LARVAL DROSOPHILA MELANOGASTER EXHIBIT GENETICALLY BASED TRADE‐OFFS AGAINST DIFFERENT SPECIES OF PARASITOIDS

Populations of Drosophila melanogaster face significant mortality risks from parasitoid wasps that use species‐specific strategies to locate and survive in hosts. We tested the hypothesis that parasitoids with different strategies select for alternative host defense characteristics and in doing so contribute to the maintenance of fitness variation and produce trade‐offs among traits. We characterized defense traits of Drosophila when exposed to parasitoids with different host searching behaviors (Aphaereta sp. and Leptopilina boulardi). We used host larvae with different natural alleles of the gene Dopa decarboxylase (Ddc), a gene controlling the production of dopamine and known to influence the immune response against parasitoids. Previous population genetic analyses indicate that our focal alleles are maintained by balancing selection. Genotypes exhibited a trade‐off between the immune response against Aphaereta sp. and the ability to avoid parasitism by L. boulardi. We also identified a trade‐off between the ability to avoid parasitism by L. boulardi and larval competitive ability as indicated by differences in foraging and feeding behavior. Genotypes differed in dopamine levels potentially explaining variation in these traits. Our results highlight the potential role of parasitoid biodiversity on host fitness variation and implicate Ddc as an antagonistic pleiotropic locus influencing larval fitness traits.     

Genetic variation in the behavioural mechanisms involved in the response of the egg parasitoid Trissolcus brochymenae to contact chemical cues left by the pest Murgantia histrionica

1. The ability of parasitoid females to perceive chemical traces left by their hosts is of utmost importance in the host location process. The behaviours involved in such ability have thus most likely been promoted by natural selection in the course of the evolutionary time. For this to happen, however, there must be significant genetic variation in natural populations on which natural selection could act.
2. Using the isofemale line method and motion analysis, we detected significant intra‐population genetic variation for several walking behaviour traits of the egg parasitoid Trissolcus brochymenae (Hymenoptera: Scelionidae) females responding to chemical traces left by its host Murgantia histrionica (Heteroptera: Pentatomidae).
3. Besides opening new avenues of research on the reproductive strategies, behaviour, and biological control potential of parasitoid wasps, these results also have implications for understanding their life‐history evolution in general.

     

Population genetic structure of two primary parasitoids of Spodoptera frugiperda (Lepidoptera), Chelonus insularis and Campoletis sonorensis (Hymenoptera): to what extent is the host plant important?

Plant chemistry can strongly influence interactions between herbivores and their natural enemies, either by providing volatile compounds that serve as foraging cues for parasitoids or predators, or by affecting the quality of herbivores as hosts or prey. Through these effects plants may influence parasitoid population genetic structure. We tested for a possible specialization on specific crop plants in Chelonus insularis and Campoletis sonorensis, two primary parasitoids of the fall armyworm, Spodoptera frugiperda. Throughout Mexico, S. frugiperda larvae were collected from their main host plants, maize and sorghum and parasitoids that emerged from the larvae were used for subsequent comparison by molecular analysis. Genetic variation at eight and 11 microsatellites were respectively assayed for C. insularis and C. sonorensis to examine isolation by distance, host plant and regional effects. Kinship analyses were also performed to assess female migration among host‐plants. The analyses showed considerable within population variation and revealed a significant regional effect. No effect of host plant on population structure of either of the two parasitoid species was found. Isolation by distance was observed at the individual level, but not at the population level. Kinship analyses revealed significantly more genetically related—or kin—individuals on the same plant species than on different plant species, suggesting that locally, mothers preferentially stay on the same plant species. Although the standard population genetics parameters showed no effect of plant species on population structure, the kinship analyses revealed that mothers exhibit plant species fidelity, which may speed up divergence if adaptation were to occur.     

Marked host specificity and lack of phylogeographic population structure of Campylobacter jejuni in wild birds

Zoonotic pathogens often infect several animal species, and gene flow among populations infecting different host species may affect the biological traits of the pathogen including host specificity, transmissibility and virulence. The bacterium Campylobacter jejuni is a widespread zoonotic multihost pathogen, which frequently causes gastroenteritis in humans. Poultry products are important transmission vehicles to humans, but the bacterium is common in other domestic and wild animals, particularly birds, which are a potential infection source. Population genetic studies of C. jejuni have mainly investigated isolates from humans and domestic animals, so to assess C. jejuni population structure more broadly and investigate host adaptation, 928 wild bird isolates from Europe and Australia were genotyped by multilocus sequencing and compared to the genotypes recovered from 1366 domestic animal and human isolates. Campylobacter jejuni populations from different wild bird species were distinct from each other and from those from domestic animals and humans, and the host species of wild bird was the major determinant of C. jejuni genotype, while geographic origin was of little importance. By comparison, C. jejuni differentiation was restricted between more phylogenetically diverse farm animals, indicating that domesticated animals may represent a novel niche for C. jejuni and thereby driving the evolution of those bacteria as they exploit this niche. Human disease is dominated by isolates from this novel domesticated animal niche.     

Inter‐oceanic variation in patterns of host‐associated divergence in a seabird ectoparasite

Aim Parasites with global distributions and wide host spectra provide excellent models for exploring the factors that drive parasite diversification. Here, we tested the relative force of host and geography in shaping population structure of a widely distributed and common ectoparasite of colonial seabirds, the tick Ixodes uriae. Location Two natural geographic replicates of the system: numerous seabird colonies of the North Pacific and North Atlantic Ocean basins. Methods Using eight microsatellite markers and tick samples from a suite of multi‐specific seabird colonies, we examined tick population structure in the North Pacific and compare patterns of diversity and structure to those in the Atlantic basin. Analyses included population genetic estimations of diversity and population differentiation, exploratory multivariate analyses, and Bayesian clustering approaches. These different analyses explicitly took into account both the geographic distance among colonies and host use by the tick. Results Overall, little geographic structure was observed among Pacific tick populations. However, host‐related genetic differentiation was evident, but was variable among host types and lower than in the North Atlantic. Main conclusions Tick population structure is concordant with the genetic structure observed in seabird host species within each ocean basin, where seabird populations tend to be less structured in the North Pacific than in the North Atlantic. Reduced tick genetic structure in the North Pacific suggests that host movement among colonies, and thus tick dispersal, is higher in this region. In addition to information on parasite diversity and gene flow, our findings raise interesting questions about the subtle ways that host behaviour, distribution and phylogeographic history shape the genetics of associated parasites across geographic landscapes.     

Host plant‐related genomic differentiation in the European cherry fruit fly,Rhagoletis cerasi

Elucidating the mechanisms and conditions facilitating the formation of biodiversity are central topics in evolutionary biology. A growing number of studies imply that divergent ecological selection may often play a critical role in speciation by counteracting the homogenising effects of gene flow. Several examples involve phytophagous insects, where divergent selection pressures associated with host plant shifts may generate reproductive isolation, promoting speciation. Here, we use ddRADseq to assess the population structure and to test for host‐related genomic differentiation in the European cherry fruit fly, Rhagoletis cerasi (L., 1758) (Diptera: Tephritidae). This tephritid is distributed throughout Europe and western Asia, and has adapted to two different genera of host plants, Prunus spp. (cherries) and Lonicera spp. (honeysuckle). Our data imply that geographic distance and geomorphic barriers serve as the primary factors shaping genetic population structure across the species range. Locally, however, flies genetically cluster according to host plant, with consistent allele frequency differences displayed by a subset of loci between Prunus and Lonicera flies across four sites surveyed in Germany and Norway. These 17 loci display significantly higher F_ST values between host plants than others. They also showed high levels of linkage disequilibrium within and between Prunus and Lonicera flies, supporting host‐related selection and reduced gene flow. Our findings support the existence of sympatric host races in R. cerasi embedded within broader patterns of geographic variation in the fly, similar to the related apple maggot, Rhagoletis pomonella, in North America.