Testing for host‐associated differentiation in two egg parasitoids of a forest herbivore

Several studies underline the importance of ecological barriers and differential selection in driving sympatric speciation. Host‐associated differentiation (HAD) has been proposed as one of the mechanisms leading to sympatric speciation. However, it is still unclear how common HAD is or which are the factors that could promote it. In particular, not much is known about HAD in predators and parasitoids of herbivorous insects. One of the characteristics postulated to pre‐dispose insects to HAD is parthenogenesis as it may favour adaptive responses to particular environments, amplifying selected gene complexes. In this study, we used amplified fragment length polymorphism (AFLP) markers to determine whether HAD is present in two parthenogenetic egg parasitoids attacking the same herbivore species – the pine processionary moth, Thaumetopoea pityocampa (Denis & Schiffermüller) (Lepidoptera: Notodontidae) – on two host Pinus species. A total of 100 loci for 59 individuals sampled in four populations of Baryscapus servadeii (Domenichini) (Hymenoptera: Eulophidae), a specialist parasitoid, and 106 loci for 117 individuals sampled in six populations of Ooencyrtus pityocampae Mercet (Hymenoptera: Encyrtidae), a generalist parasitoid, were analysed. Levels of genetic differentiation were also assessed with an outlier analysis, checking for alleles associated to host plants. No evidence of HAD was detected in any of the two parasitoid species. We hypothesize that both the lack of strict parthenogenetic reproduction and the ectophagous nature of the insect host could explain the absence of HAD. The genetic variation observed in the generalist parasitoid responded to a pattern of local adaptation, whereas no relationship with either host or geography was found in the specialist parasitoid.     

Spatial and host related genomic variation in partially sympatric cactophagous moth species

Surveys of patterns of genetic variation in natural sympatric and allopatric populations of recently diverged species are necessary to understand the processes driving intra- and interspecific diversification. The South American moths Cactoblastis cactorum, Cactoblastis doddi and Cactoblastis bucyrus are specialized in the use of cacti as host plants. These species have partially different geographic ranges and differ in patterns of host plant use. However, there are areas that overlap, particularly, in northwestern Argentina, where they are sympatric. Using a combination of genome-wide SNPs and mitochondrial data we assessed intra and interspecific genetic variation and investigated the relative roles of geography and host plants on genetic divergence. We also searched for genetic footprints of hybridization between species. We identified three well delimited species and detected signs of hybridization in the area of sympatry. Our results supported a hypothetical scenario of allopatric speciation in the generalist C. cactorum and genetic interchange during secondary geographic contact with the pair of specialists C. bucyrus and C. doddi that probably speciated sympatrically. In both cases, adaptation to new host plants probably played an important role in speciation. The results also suggested the interplay of geography and host plant use as drivers of divergence and limiting gene flow at intra and interspecific levels.     

Geographic pattern of host‐associated differentiation in the cotton fleahopper,Pseudatomoscelis seriatus

Host‐associated differentiation (HAD) is the occurrence of genetically distinct, host‐associated lineages. Most of the cases of HAD in phytophagous insects have been documented in specialist insects inhabiting feral ecosystems or in generalist parthenogens in agroecosystems. Herein we report HAD in the cotton fleahopper, Pseudatomoscelis seriatus (Reuter) (Hemiptera: Miridae), a native, generalist, non‐parthenogenetic insect feeding on native wild hosts [horsemint, Monarda punctata L. (Lamiaceae) and woolly croton, Croton capitatus Michx. (Euphorbiaceae)] and on cotton [Gossypium hirsutum L. (Malvaceae)] in the USA. Examination of genome‐wide genetic variation with AFLP markers and Bayesian analyses of P. seriatus associated with three different host plant species at five locations in Texas revealed a geographic pattern of HAD. The geographic pattern of HAD corresponded with differences in precipitation among the locations studied. In three locations, two distinct lineages of P. seriatus were found in association with horsemint and cotton/woolly croton, whereas in two other locations, populations associated with the different host plants studied were panmictic. We suggest that precipitation differences among locations translate into heterogeneity in vegetation distribution, composition, and phenology, which altogether may contribute to the observed geographic pattern of HAD.     

Spatial and genetic structure of host‐associated differentiation in the parasitoid Copidosoma gelechiae

Host‐associated differentiation (HAD) appears to be an important driver of diversification in the hyperdiverse phytophagous and parasitoid insects. The gallmaking moth Gnorimoschema gallaesolidaginis has undergone HAD on two sympatric goldenrods (Solidago), and HAD has also been documented in its parasitoid Copidosoma gelechiae, with the intriguing suggestion that differentiation has proceeded independently in multiple populations. We tested this suggestion with analysis of Amplified Fragment Length Polymorphism (AFLP) markers for C. gelechiae collections from the midwestern and northeastern United States and eastern Canada. AFLP data were consistent with the existence of HAD, with between‐host F_ST significant before Bonferroni correction in two of seven sympatric populations. amova analysis strongly rejected a model of HAD with a single historical origin, and thus supported the repeated‐HAD hypothesis. Copidosoma gelechiae shows significant host‐associated divergence at a number of allozyme loci ( Stireman et al., 2006 ), but only weak evidence via AFLPs for genome‐wide differentiation, suggesting that this species is at a very early stage of HAD.     

Host‐associated differentiation in a highly polyphagous,sexually reproducing insect herbivore

Insect herbivores may undergo genetic divergence on their host plants through host‐associated differentiation (HAD). Much of what we know about HAD involves insect species with narrow host ranges (i.e., specialists) that spend part or all their life cycle inside their hosts, and/or reproduce asexually (e.g., parthenogenetic insects), all of which are thought to facilitate HAD. However, sexually reproducing polyphagous insects can also exhibit HAD. Few sexually reproducing insects have been tested for HAD, and when they have insects from only a handful of potential host‐plant populations have been tested, making it difficult to predict how common HAD is when one considers the entire species' host range. This question is particularly relevant when considering insect pests, as host‐associated populations may differ in traits relevant to their control. Here, we tested for HAD in a cotton (Gossypium hirsutum) pest, the cotton fleahopper (CFH) (Pseudatomoscelis seriatus), a sexually reproducing, highly polyphagous hemipteran insect. A previous study detected one incidence of HAD among three of its host plants. We used Amplified fragment length polymorphism (AFLP) markers to assess HAD in CFH collected from an expanded array of 13 host‐plant species belonging to seven families. Overall, four genetically distinct populations were found. One genetically distinct genotype was exclusively associated with one of the host‐plant species while the other three were observed across more than one host‐plant species. The relatively low degree of HAD in CFH compared to the pea aphid, another hemipteran insect, stresses the likely importance of sexual recombination as a factor increasing the likelihood of HAD.     

Rapid evolution and specialization following host colonization in a yucca moth

Host shifts followed by specialization can result in sympatric genetic differentiation, and may have fuelled the diversification of phytophagous insects. This study examines a recent colonization of a non‐native host by Prodoxus quinquepunctellus (Lepidoptera: Prodoxidae). Allozyme differentiation was detected among different host feeding populations, yet was nearly absent among similar host feeding populations in sympatry. Geographical patterns of allozyme variation showed a much higher level of population structure among populations feeding on the derived host. Conversely, mtDNA haplotype frequencies were nearly homogeneous in the derived populations compared to the ancestral populations, suggesting a bottleneck and/or rapid fixation of haplotypes following host colonization. Moth emergence coincided with host plant flowering, and phenological differences between host species translated into allochronic isolation between populations feeding on different hosts. Derived moth populations also differed significantly in three ovipositor characters from ancestral populations. These findings suggest rapid host‐specific genetic differentiation, and specialization of moth emergence time and ovipositor morphology following host colonization.     

Host and geography together drive early adaptive radiation of Hawaiian planthoppers

The interactions between insects and their plant host have been implicated in driving diversification of both players. Early arguments highlighted the role of ecological opportunity, with the idea that insects “escape and radiate” on new hosts, with subsequent hypotheses focusing on the interplay between host shifting and host tracking, coupled with isolation and fusion, in generating diversity. Because it is rarely possible to capture the initial stages of diversification, it is particularly difficult to ascertain the relative roles of geographic isolation versus host shifts in initiating the process. The current study examines genetic diversity between populations and hosts within a single species of endemic Hawaiian planthopper, Nesosydne umbratica (Hemiptera, Delphacidae). Given that the species was known as a host generalist occupying unrelated hosts, Clermontia (Campanulaceae) and Pipturus (Urticaceae), we set out to determine the relative importance of geography and host in structuring populations in the early stages of differentiation on the youngest islands of the Hawaiian chain. Results from extensive exon capture data showed that N. umbratica is highly structured, both by geography, with discrete populations on each volcano, and by host plant, with parallel radiations on Clermontia and Pipturus leading to extensive co‐occurrence. The marked genetic structure suggests that populations can readily become established on novel hosts provided opportunity; subsequent adaptation allows monopolization of the new host. The results support the role of geographic isolation in structuring populations and with host shifts occurring as discrete events that facilitate subsequent parallel geographic range expansion.     

Behavioural,ecological and genetic evidence confirm the occurrence of host‐associated differentiation in goldenrod gall‐midges

Host‐associated differentiation (HAD) is considered a step towards ecological speciation and an important mechanism promoting diversification in phytophagous insects. Although the number of documented cases of HAD is increasing, these still represent only a small fraction of species and feeding guilds among phytophagous insects, and most reports are based on a single type of evidence. Here we employ a comprehensive approach to present behavioural, morphological, ecological and genetic evidence for the occurrence of HAD in the gall midge Dasineura folliculi (Diptera: Cecidomyiidae) on two sympatric species of goldenrods (Solidago rugosa and S. gigantea). Controlled experiments revealed assortative mating and strong oviposition fidelity for the natal‐host species. Analysis of mitochondrial DNA showed an amount of genetic divergence between the two host‐associated populations compatible with cryptic species rather than host races. Lower levels of within‐host genetic divergence, gall development and natural‐enemy attack in the S. gigantea population suggest this is the derived host.     

Host‐associated genetic differentiation in pecan leaf phylloxera

Host‐associated differentiation (HAD) is the formation of genetically distinct host‐associated populations. One of the genotypic signatures of HAD is that populations exhibit stronger differentiation by host‐plant species than by geographic isolation. HAD, as a mechanism promoting ecological speciation, has been invoked to explain phytophagous insect diversity. Two traits proposed to promote HAD are endophagy and parthenogenesis. Using amplified fragment length polymorphisms (AFLPs), we tested for the presence of HAD in pecan leaf phylloxera, Phylloxera notabilis Pergande (Hemiptera: Phylloxeridae), an endophagous, gall inducing, and cyclically parthenogenetic insect on sympatric pecan and water hickory at a geographic mesoscale. This species shows strong HAD. Whereas the effect of collecting site was significant, accounting for 7.3% of molecular variation, host‐plant species identity accounted for 63.5%. In addition, a choice test indicated that pecan leaf phylloxera originating from water hickory showed weak but significant preference for leaflets of the natal host, whereas pecan leaf phylloxera originating from pecan did not. This is the first such study of a species of arboreal Phylloxeridae, a poorly known insect group. This is also the first endophage and the second parthenogen shared by these two hickory species to show evidence of HAD. This hickory system could be a good parthenogen‐rich counterpoint to the goldenrod system in the study of HAD in insect communities.     

Testing host‐associated differentiation in a quasi‐endophage and a parthenogen on native trees

Host‐associated differentiation (HAD) is the formation of genetically divergent host‐associated sub‐populations. Evidence of HAD has been reported for multiple insect herbivores to date, but published studies testing more than one herbivore for any given host‐plant species pair is limited to herbivores on goldenrods. This limits the number of pair‐wise comparisons that can be made about insect life‐history traits that might facilitate or inhibit host‐race development in general. Two traits previously proposed to facilitate HAD include endophagy and parthenogenesis. We tested for HAD in two herbivores, a quasi‐endophagous caterpillar and a parthenogenetic aphid, feeding on two closely related species of hickories. We found that the quasi‐endophage is panmictic, whereas the parthenogen exhibits HAD on their sympatric host plants, pecan and water hickory, at a geographic mesoscale. This is an important first step in the characterization of HAD in multiple insect herbivores using North American hickories, a host‐plant system with many shared parthenogens.     

Host plant associations and geography interact to shape diversification in a specialist insect herbivore

Disentangling the processes underlying geographic and environmental patterns of biodiversity challenges biologists as such patterns emerge from eco‐evolutionary processes confounded by spatial autocorrelation among sample units. The herbivorous insect, Belonocnema treatae (Hymenoptera: Cynipidae), exhibits regional specialization on three plant species whose geographic distributions range from sympatry through allopatry across the southern United States. Using range‐wide sampling spanning the geographic ranges of the three host plants and genotyping‐by‐sequencing of 1,217 individuals, we tested whether this insect herbivore exhibited host plant‐associated genomic differentiation while controlling for spatial autocorrelation among the 58 sample sites. Population genomic structure based on 40,699 SNPs was evaluated using the hierarchical Bayesian model entropy to assign individuals to genetic clusters and estimate admixture proportions. To control for spatial autocorrelation, distance‐based Moran's eigenvector mapping was used to construct regression variables summarizing spatial structure inherent among sample sites. Distance‐based redundancy analysis (dbRDA) incorporating the spatial variables was then applied to partition host plant‐associated differentiation (HAD) from spatial autocorrelation. By combining entropy and dbRDA to analyse SNP data, we unveiled a complex mosaic of highly structured differentiation within and among gall‐former populations finding evidence that geography, HAD and spatial autocorrelation all play significant roles in explaining patterns of genomic differentiation in B. treatae. While dbRDA confirmed host association as a significant predictor of patterns of genomic variation, spatial autocorrelation among sites explained the largest proportion of variation. Our results demonstrate the value of combining dbRDA with hierarchical structural analyses to partition spatial/environmental patterns of genomic variation.     

Geography is more important than host plant use for the population genetic structure of a generalist insect herbivore

Population divergence can occur due to mechanisms associated with geographic isolation and/or due to selection associated with different ecological niches. Much of the evidence for selection‐driven speciation has come from studies of specialist insect herbivores that use different host plant species; however, the influence of host plant use on population divergence of generalist herbivores remains poorly understood. We tested how diet breadth, host plant species and geographic distance influence population divergence of the fall webworm (Hyphantria cunea; FW). FW is a broadly distributed, extreme generalist herbivore consisting of two morphotypes that have been argued to represent two different species: black‐headed and red‐headed. We characterized the differentiation of FW populations at two geographic scales. We first analysed the influence of host plant and geographic distance on genetic divergence across a broad continental scale for both colour types. We further analysed the influence of host plant, diet breadth and geographic distance on divergence at a finer geographic scale focusing on red‐headed FW in Colorado. We found clear genetic and morphological distinction between red‐ and black‐headed FW, and Colorado FW formed a genetic cluster distinct from other locations. Although both geographic distance and host plant use were correlated with genetic distance, geographic distance accounted for up to 3× more variation in genetic distance than did host plant use. As a rare study investigating the genetic structure of a widespread generalist herbivore over a broad geographic range (up to 3,000 km), our study supports a strong role for geographic isolation in divergence in this system.     

Divergence in host use ability of a marine herbivore from two habitat types

We studied ecological divergence of host use ability in a generalist marine herbivore living in two distinct host plant assemblages. We collected Idotea balthica isopods from three populations dominated by the brown alga Fucus vesiculosus and three dominated by the seagrass Zostera marina. In two reciprocal common garden feeding experiments for adult and laboratory‐born juvenile isopods, we found that isopods from both assemblages performed better with their sympatric dominant host species than did isopods allopatric to this host. This indicates parallel divergence of populations according to the sympatric host plant assemblage. Furthermore, initial body size and body size‐dependent mortality differed between populations from the two assemblages. In nature, this may result in lower fitness of immigrants compared with that of residents and consequently reinforce divergence of the populations. Finally, we discuss how phenotypic plasticity and maternal and random effects may associate with the results.     

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.     

Predicting the outcomes of a tri‐trophic interaction between an indigenous parasitoid and an exotic herbivorous pest and its host plants

Understanding the novel ecological interactions that result from biological invasions is a critical issue in modern ecology and evolution as well as pest management. Introduced herbivorous insects may interact with native plants and indigenous natural enemies, creating novel tri‐trophic interactions. To help predict the potential outcomes of novel interactions, we investigated the behavioural and physiological responses of an indigenous generalist parasitoid (Habrobracon gelechiae) to an introduced generalist herbivore (the light brown apple moth, Epiphyas postvittana) and its new host plants in California. We first examined the parasitoid's host location and acceptance on a range of nine common host plants of the moth representing distinctly different geographic origins and morphologies (to examine the effect of a known toxic plant on the parasitoid's performance, an additional toxic plant species was also tested that the moth consumes in the laboratory but does not naturally attack). The parasitoid was able to locate the host larvae on all plants equally well, although clutch size was affected by host plant. We then determined fitness of the moth and the parasitoid on four representative plants. The moth larvae suffered higher mortality and a slower developmental rate on the known toxic plant than on the other three plants, but the parasitoid's fitness correlates did not differ between the host food plants. These results show a high level of plasticity in the indigenous generalist parasitoid in its ability to exploit the exotic host on a wide range of host plants, generating an invasion‐driven novel tri‐trophic interaction.     

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.     

Molecular evidence of host-associated genetic divergence in the holly leafminer Phytomyza glabricola (Diptera: Agromyzidae): apparent discordance among marker systems

Host races play a central part in understanding the role of host plant mediated divergence and speciation of phytophagous insects. Of greatest interest are host-associated populations that have recently diverged; however, finding genetic evidence for very recent divergences is difficult because initially only a few loci are expected to evolve diagnostic differences. The holly leafminer Phytomyza glabricola feeds on two hollies, Ilex glabra and I. coriacea, that are broadly sympatric throughout most of their ranges. The leafminer is often present on both host plants and exhibits a dramatic life history difference on the two hosts, suggesting that host races may be present. We collected 1393 bp of mitochondrial cytochrome oxidase I (COI) sequence and amplified fragment length polymorphism (AFLP) data (45 polymorphic bands) from sympatric populations of flies reared from the two hosts. Phylogenetic and frequency analysis of mitochondrial COI sequence data uncovered considerable variation but no structuring by the host plant, and only limited differentiation among geographical locations. In contrast, analysis of AFLP frequency data found a significant effect with host plant, and a much smaller effect with geographical location. Likewise, neighbour-joining analysis of AFLP data resulted in clustering by host plant. The AFLP data indicate that P. glabricola is most likely comprised of two host races. Because there were no fixed differences in mitochondrial or AFLP data, this host-associated divergence is likely to have occurred very recently. P. glabricola therefore provides a new sympatric system for exploring the role of geography and ecological specialization in the speciation of phytophagous insects.     

GENETIC AND GEOGRAPHIC VARIATION IN REJECTION BEHAVIOR OF CUCKOO EGGS BY EUROPEAN MAGPIE POPULATIONS: AN EXPERIMENTAL TEST OF REJECTER-GENE FLOW

Host responses toward brood parasitism have been shown to differ among populations depending on the duration of sympatry between host and parasite, although populations not currently parasitized show rejection behavior against parasitic eggs. The persistence of rejection behavior in unparasitized host populations and rapid increases of rejection rate in parasitized ones have sometimes been explained as the result of gene flow of rejecter genes from sympatry to allopatry (rejecter-gene flow hypothesis). We present data on the rejection behavior of magpies (Pica pica) the main European host of the great spotted cuckoo (Clamator glandarius), in 15 populations (nine sympatric six allopatric) across their distribution range in Europe. Rejection rates of mimetic and nonmimetic model eggs were significantly higher in sympatric than in allopatric magpie populations, although differences in rejection rate of both mimetic and nonmimetic model eggs between magpie populations were significantly correlated even after controlling tor phylogenetic effects, with differences between sympatric and allopatric magpie populations being larger for mimetic than for nonmimetic model eggs. Differences in rejection of mimetic model eggs were related to both genetic and geographic distances between populations, but differences in rejection rate of nonmimetic model eggs were unrelated to these distances. However, when comparing only sympatric populations, differences in rejection rate of both mimetic and nonmimetic model eggs were related to geographic distances. A multiple autocorrelation analysis revealed that differences among populations in rejection rates of mimetic model eggs had a strong geographic component whereas the main component of rejection rate of nonmimetic model eggs was genetic rather than geographic. These results support the rejecter-gene flow hypothesis. We discuss differences in rejection rates of mimetic and nonmimetic model eggs that suggest the egg-recognition ability of the host is genetically based, but is affected by a learning process for fine tuning of recognition.     

HOST‐ASSOCIATED GENETIC DIFFERENTIATION IN PHYTOPHAGOUS INSECTS: GENERAL PHENOMENON OR ISOLATED EXCEPTIONS? EVIDENCE FROM A GOLDENROD‐INSECT COMMUNITY

There is growing awareness of the importance of natural selection in driving genetic divergence and speciation, and several of the most apparent cases of this ecological speciation are provided by the existence of genetically distinct host forms in phytophagous insects. Such examples of host-associated differentiation (HAD) have become increasingly documented, and the implications of this phenomenon for the diversification of insects are becoming widely appreciated. However, instances of HAD remain rare relative to insect diversity and are sparsely distributed both ecologically and taxonomically. We sought to assess the frequency of HAD in a model herbivore community by examining genetic divergence in a variety of herbivores that feed on two closely related and broadly sympatric species of goldenrod (Solidago altissima and S. gigantea). Using mitochondrial DNA and allozyme data, in conjunction with previously published studies, we found that four of nine herbivores exhibited evidence of HAD, including possible host races or cryptic species. Using a range of reasonable substitution rate estimates for cytochrome oxidase I mitochondrial DNA, we found that HAD appears to have proceeded asynchronously across taxa. This pattern, along with the broadly sympatric distribution of host plants and the specialized life histories of the phytophagous insects, is consistent with sympatric divergence in some or all of these taxa. Although further behavioral and ecological study is needed, our survey of HAD in a community of herbivores indicates that ecological (perhaps sympatric) speciation may have been responsible for generating a significant fraction of the extant diversity of phytophagous insects.     

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.