Host Race Formation Research Articles

Host‐specific morphologies but no host races in the commensal bivalveNeaeromya rugifera

AbstractSpeciation by host shift is one of the explicit models of ecological speciation. A prerequisite of this model is the formation of host races (sympatric populations that show host‐specific genetic structuring and phenotypes). Many members of the diverse marine bivalve superfamilyGaleommatoidea have obligate commensal relationships with invertebrate hosts. Some species have the ability to occupy multiple host species, thereby providing potential opportunities to test for the formation of host races. TheNortheastPacific galeommatoideanNeaeromya rugiferaattaches to two strikingly different hosts: the blue mud shrimpUpogebia pugettensisand the polychaete sea mouseAphroditaspp. We tested if this host difference has resulted in the formation of host races using shell morphologies and genetic markers. We found that populations from different hosts differ significantly in shell morphology. However, based on mitochondrial makers,N. rugiferashowed no distinct host‐specific genetic structuring, indicating the existence of a panmictic population. We conclude that the host‐specific morphologies these clams exhibit may reflect ecophenotypic plasticity rather than the existence of host races, but this needs to be corroborated with additional genetic data and larger sample sizes. The pronounced conchological variation withinN. rugiferacalls for further investigation of its taxonomic relationship with its poorly studied, but morphologically similar, sympatric congenerNeaeromya compressa.

Environmental interactions during host race formation: host fruit environment moderates a seasonal shift in phenology in host races of Rhagoletis pomonella

Summary1. Host race formation is a common form of ecological speciation during which new populations that exploit a novel host (e.g. nutritional resource) experience divergent natural selection, causing adaptive divergence from the ancestral population. Typically, multiple selection pressures drive this divergence, suggesting that interactions among environmental effects may be critical during the speciation process.2. Host‐race‐forming phytophagous insects often experience divergent natural selection imposed by seasonality and nutritional environment, two factors likely to interact through their effects on growth and life‐history timing. We tested for the presence of such an interaction in the apple maggot fly, Rhagoletis pomonella, asking whether nutritional effects interact with seasonality to determine overwintering success.3. We find evidence for such an interaction, wherein feeding on a novel host fruit actually mitigates the negative effects of novel seasonality. Feeding on apple (novel) compared to hawthorn (ancestral) fruit conferred greater fly lipid reserves the overwintering, diapause stage. Earlier, seasonal emergence characteristic of the apple host race imposes an overwintering survival cost. But feeding on apples offsets this cost, which confers greater pupal lipid reserves and consequently larger adult body size post‐winter. Seasonality and host fruit interact to determine fly lipid reserves, and host race differences in lipid content are maximized under the most stressful conditions typically experienced by the apple host race. F1 rearing and genetic association tests revealed no evidence for genetically based divergence in lipid content, suggesting that differences in lipid storage among the fly host races are driven primarily by the host fruit environment.4. Our results suggest that interactions between seasonality and host plant environment shape natural selection and therefore influence adaptive divergence during host race formation.

Geographic variation in fruit volatiles emitted by the hawthorn Crataegus mollis and its consequences for host race formation in the apple maggot fly, Rhagoletis pomonella

AbstractThe shift of the apple (AP) maggot fly, Rhagoletis pomonella (Walsh) (Diptera: Tephritidae), from its ancestral host downy hawthorn, Crataegus mollis (DH) (Torr. & A. Gray) Scheele, to introduced domesticated AP, Malus domestica Borkh. (both Rosaceae), is a model for ecological divergence and incipient sympatric speciation with gene flow. However, a portion of the variation contributing to the sympatric host shift from DH to AP appears to have a different biogeographic history, pre‐dating the shift. One potential source of standing variation may trace to a number of different native hawthorn species infested by R. pomonella in the southern USA, where the AP‐attacking race is absent. Herein, we investigate this possibility for the southern red hawthorn (SR) endemic to Texas, Crataegus mollis var. texana (Buckl.), which has been described as a member of the Molles series that includes the more northern distributed DH. We report results from chemical analyses of host fruit volatiles, fly behavioural responses to synthetic fruit blends, and microsatellite surveys of fly populations, implying that R. pomonella infesting SR may behaviourally and genetically represent a native host race differing from the DH‐infesting fly. No fly reared from SR responded to AP fruit volatiles in flight tunnel assays. However, coupled gas chromatographic‐electroantennographic detection (GC‐EAD) profiles for SR fruit contain all five of the component esters that comprise the standard AP volatile blend inducing behavioural orientation for AP‐infesting flies, compounds that appear to be largely missing from volatile profiles for DH fruit. Thus, SR‐infesting flies do not represent a source for a preassembled AP‐accepting phenotype. However, they may help explain why the ancestral DH race that shifted to AP in the northeastern USA had the ability to recognize AP fruit esters, potentially enabling the shift to AP. Our results highlight how categorizing speciation into different geographic modes may not adequately describe the evolutionary origins of important genetic variation fuelling adaptive radiation and the genesis of new biodiversity.

High divergence levels of Panonychus citri populations on Rutaceae and Oleaceae as indicated by internal transcribed spacer 1 (ITS1) sequences

The citrus red mite, Panonychus citri (McGregor) (Acari: Tetranychidae), has a worldwide distribution and is the most important citrus pest in many countries, including Japan, Spain and China. Apart from citrus, P. citri also feeds on other woody plants such as pear, peach and holly. To characterize the population genetic structure of citrus red mite in association with their host plants, the gene flow (Nm) and genetic differentiation among sympatric populations of P. citri (F ST estimates) on different host plants were estimated using the ribosomal internal transcribed spacer 1 (ITS1) sequence. The results indicated that there were high levels of gene flow (Nm = infinite) among the populations from the same host genera and the same host family. Populations of P. citri displayed genetic differentiation and low levels of gene flow (Nm < 1) between the populations collected from Rutaceae and Oleaceae host plants. While the haplotype network presented a parallel pattern, there was no one central common haplotype. Pairwise F ST indicated that in China, P. citri has differentiated into two groups, one of which was associated with sweet olive and the other with other host plants. The result of cytochrome oxidase subunit I analysis also supported this result, suggesting the possibility of sympatric host race formation in this species.

Developmental trajectories of gene expression reveal candidates for diapause termination: a key life-history transition in the apple maggot flyRhagoletis pomonella

The timing of dormancy is a rapidly evolving life-history trait playing a crucial role in the synchronization of seasonal life cycles and adaptation to environmental change. But the physiological mechanisms regulating dormancy in animals remain poorly understood. In insects, dormancy (diapause) is a developmentally dynamic state, and the mechanisms that control diapause transitions affect seasonal timing. Here we used microarrays to examine patterns of gene expression during dormancy termination: a crucial life-history transition in the apple maggot fly Rhagoletis pomonella (Walsh). This species is a model system for host race formation and ecological speciation via changes in diapause regulation of seasonality. Our goal was to pinpoint the timing of the transition from diapause to post-diapause development and to identify candidate genes and pathways for regulation of diapause termination. Samples were taken at six metabolically defined developmental landmarks, and time-series analysis suggests that release from metabolic depression coincides with preparation for or resumption of active cell cycling and morphogenesis, defining the 'end' of diapause. However, marked changes in expression, including members of pathways such as Wnt and TOR signaling, also occur prior to the metabolic rate increase, electing these pathways as candidates for early regulation of diapause termination. We discuss these results with respect to generalities in insect diapause physiology and to our long-term goal of identifying mechanisms of diapause adaptation in the Rhagoletis system.

Identification of Fruit Volatiles from Green Hawthorn (Crataegus Viridis) and Blueberry Hawthorn (Crataegus Brachyacantha) Host Plants Attractive to Different Phenotypes of Rhagoletis Pomonella Flies in the Southern United States

The apple maggot fly, Rhagoletis pomonella, infests several hawthorn species in the southern USA. In a companion paper, we showed that R. pomonella flies infesting two different mayhaw species (Crataegus opaca and C. aestivalis) can discriminate between volatile blends developed for each host fruit, and that these blends are different from previously constructed blends for northern fly populations that infest domestic apple (Malus domestica), downy hawthorn (Crataegus mollis), and flowering dogwood (Cornus florida). Here, we show by using coupled gas chromatography and electroantennographic detection (GC-EAD), gas chromatography with mass spectrometry (GC-MS), and flight tunnel bioassays, that two additional southern hawthorn fly populations infesting C. viridis (green hawthorn) and C. brachyacantha (blueberry hawthorn) also can discriminate between volatile blends for each host fruit type. A 9-component blend was developed for C. viridis (3-methylbutan-1-ol [5%], butyl butanoate [19.5%], propyl hexanoate [1.5%], butyl hexanoate [24%], hexyl butanoate [24%], pentyl hexanoate [2.5%], 1-octen-3-ol [0.5%], pentyl butanoate [2.5%], and (3E)-4,8-dimethyl-1,3,7-nonatriene (DMNT) [20.5%]) and an 8-component blend for C. brachyacantha (3-methylbutan-1-ol [0.6%], butyl acetate [50%], pentyl acetate [3.5%], butyl butanoate [9%], butyl hexanoate [16.8%], hexyl butanoate [16.8%], 1-octen-3-ol [0.3%], and pentyl butanoate [3%]). Crataegus viridis and C. brachyacantha-origin flies showed significantly higher levels of upwind oriented flight to their natal blend in flight tunnel assays compared to the alternate, non-natal blend and previously developed northern host plant blends. The presence of DMNT in C. viridis and butyl acetate in C. brachyacantha appeared to be largely responsible for driving the differential response. This sharp behavioral distinction underscores the diversity of odor response phenotypes in the southern USA, points to possible host race formation in these populations, and despite the presence of several apple volatiles in both blends, argues against a functional apple race existing on southern host plants prior to the introduction of apple to North America.

Identification of Host Fruit Volatiles from Three Mayhaw Species (Crataegus Series Aestivales) Attractive to Mayhaw-Origin Rhagoletis pomonella Flies in the Southern United States

The apple maggot fly, Rhagoletis pomonella, infests several hawthorn species in the southern USA. Here, we tested the hypothesis that these populations could serve as reservoirs for fruit odor discrimination behaviors facilitating sympatric host race formation and speciation, specifically the recent shift from downy hawthorn (Crataegus mollis) to domestic apple (Malus domestica) in the northern USA. Coupled gas chromatography and electroantennographic detection (GC-EAD), gas chromatography with mass spectrometry (GC-MS), and flight tunnel bioassays were used to identify the behaviorally active natal fruit volatile blends for three of the five major southern hawthorns: C. opaca (western mayhaw), C. aestivalis (eastern mayhaw), and C. rufula (a possible hybrid between C. opaca and C. aestivalis). A 6-component blend was developed for C. opaca (3-methylbutan-1-ol [44%], pentyl acetate [6%], butyl butanoate [6%], propyl hexanoate [6%], butyl hexanoate [26%], and hexyl butanoate [12%]); an 8-component blend for C. aestivalis (3-methylbutan-1-ol [2%], butyl acetate [47%], pentyl acetate [2%], butyl butanoate [12%], propyl hexanoate [1%], butyl hexanoate [25%], hexyl butanoate [9%], and pentyl hexanoate [2%]); and a 9-component blend for C. rufula (3-methylbutan-1-ol [1%], butyl acetate [57%], 3-methylbutyl acetate [3%], butyl butanoate [5%], propyl hexanoate [1%], hexyl propionate [1%], butyl hexanoate [23%], hexyl butanoate [6%], and pentyl hexanoate [3%]). Crataegus aestivalis and C. opaca-origin flies showed significantly higher levels of upwind directed flight to their natal blend in flight tunnel assays compared to the non-natal blend and previously developed apple, northern downy hawthorn, and flowering dogwood blends. Eastern and western mayhaw flies also were tested to the C. rufula blend, with eastern flies displaying higher levels of upwind flight compared with the western flies, likely due to the presence of butyl acetate in the C. aestivalis and C. rufula blends, an agonist compound for eastern mayhaw-origin flies, but a behavioral antagonist for western flies. The results discount the possibility that the apple fly was "pre-assembled" and originated via a recent introduction of southern mayhaw flies predisposed to accepting apple. Instead, the findings are consistent with the possibility of southern mayhaw-infesting fly host races. However, mayhaw fruits do emit several volatiles found in apple. It is, therefore, possible that the ability of the fly to evolve a preference for apple volatiles, although not the entire blend, stemmed, in part, from standing variation related to the presence of these compounds in southern mayhaw fruit.

No evidence for host specialization or host-race formation in the European bitterling (Rhodeus amarus), a fish that parasitizes freshwater mussels

Coevolutionary relationships between parasites and hosts can elevate the rate of evolutionary changes owing to reciprocal adaptations between coevolving partners. Such relationships can result in the evolution of host specificity. Recent methodological advances have permitted the recognition of cryptic lineages, with important consequences for our understanding of biological diversity. We used the European bitterling (Rhodeus amarus), a freshwater fish that parasitizes unionid mussels, to investigate host specialization across regions of recent and ancient sympatry between coevolving partners. We combined genetic data (12 microsatellite and 2 mitochondrial markers) from five populations with experimental data for possible mechanisms of host species recognition (imprinting and conditioning). We found no strong evidence for the existence of cryptic lineages in R.amarus, though a small proportion of variation among individuals in an area of recent bitterling-mussel association was statistically significant in explaining host specificity. No other measures supported the existence of host-specific lineages. Behavioural data revealed a weak effect of conditioning that biased behavioural preferences towards specific host species. Host imprinting had no effect on oviposition behaviour. Overall, we established that populations of R.amarus show limited potential for specialization, manifested as weak effects of host conditioning and genetic within-population structure. Rhodeus amarus is the only species of mussel-parasitizing fish in Europe, which contrasts with the species-rich communities of bitterling in eastern Asia where several host-specific bitterling occur. We discuss costs and constraints on the evolution of host-specific lineages in our study system and more generally.

Multiple Wolbachia infections in Rhagoletis pomonella

Rhagoletis pomonella Walsh (Diptera: Tephritidae) is a model species for sympatric speciation through host race formation on apple and hawthorn. The bacterial endosymbiont Wolbachia, a manipulator of arthropod reproduction, has been considered to contribute to speciation in several species. A potential role of Wolbachia in sympatric speciation of R. pomonella remains to be tested despite an earlier detection by PCR. In this study, we isolated Wolbachia from R. pomonella individuals from both host species using multi-locus sequence typing (MLST) and the surface protein wsp. By cloning and sequencing of 311 plasmids, we found sequence types of at least four wPom strains. A complete MLST profile was obtained only for wPom1, whereas MLST loci of the other putative strains were difficult to assign because of multiple infections and low sample numbers. wPom1 occurs in both host races, whereas different sequence types were found at low frequencies only in apple-infesting R. pomonella. This warrants further investigation as it cannot be excluded that Wolbachia plays a part in this model of sympatric speciation.

Experimental evidence of host race formation inMitourabutterflies (Lepidoptera: Lycaenidae)

A population of herbivorous insects that shifts to a novel host can experience selection pressures that result in adaptation to the new resource. Host race formation, considered an early stage of the speciation process, may result. The current study investigates host shifts and variation in traits potentially involved in the evolution of reproductive isolation among populations of the juniper hairstreak butterfly,Mitoura gryneus. Mitouraare closely associated with their host trees (Cupressaceae) and exhibit host plant fidelity: in addition to larval development and oviposition, host trees support male leks and mating. Female oviposition preference for the natal host, and differential fitness of larvae when reared on natal versus alternate hosts, are indications that specialization and local adaptation to the natal host plant are occurring. Populations with single host plant associations (Juniperus ashei,J. pinchotiiandJ. virginiana) as well as populations with multiple hosts (bothJ. asheiandJ. pinchotii) were examined. Concordance between female preference and larval performance was found forJ. ashei‐associated populations. Population‐level variation in the patterns of female preference and larval performance, both within and among host associations, may reflect differences in the timing and direction of colonization of hosts. For a single nominal species that otherwise exhibits no morphological or phenological differences, the experimental assessment of specialization and host fidelity inM. gryneusprovides strong support for the hypothesis of ongoing host race formation in these butterflies.

Expressed Sequence Tags from Cephalic Chemosensory Organs of the Northern Walnut Husk Fly,Rhagoletis suavis, Including a Putative Canonical Odorant Receptor

Rhagoletis fruit flies are important both as major agricultural pests and as model organisms for the study of adaptation to new host plants and host race formation. Response to fruit odor plays a critical role in such adaptation. To better understand olfaction in Rhagoletis, an expressed sequence tag (EST) study was carried out on the antennae and maxillary palps of Rhagoletis suavis (Loew) (Diptera: Tephritidae), a common pest of walnuts in eastern United States. After cDNA cloning and sequencing, 544 ESTs were annotated. Of these, 66% had an open reading frame and could be matched to a previously sequenced gene. Based on BLAST sequence homology, 9% (49 of 544 sequences) were nuclear genes potentially involved in olfaction. The most significant finding is a putative odorant receptor (OR), RSOr1, that is homologous to Drosophila melanogaster Or49a and Or85f. This is the first tephritid OR discovered that might recognize a specific odorant. Other olfactory genes recovered included odorant binding proteins, chemosensory proteins, and putative odorant degrading enzymes.

A test of the sympatric host race formation hypothesis in Neodiprion (Hymenoptera: Diprionidae)

Theory suggests that sympatric speciation is possible; however, its prevalence in nature remains unknown. Because Neodiprion sawflies are host specialists and mate on their hosts, sympatric speciation via host shifts may be common in this genus. Here, we test this hypothesis using near-complete taxonomic sampling of a species group, comprehensive geographical and ecological data, and multiple comparative methods. Host-use data suggest that host shifts contributed to the evolution of reproductive isolation in Neodiprion and previous work has shown that gene flow accompanied divergence. However, geographical data provide surprisingly little support for the hypothesis that host shifts occurred in sympatry. While these data do not rule out sympatric host race formation in Neodiprion, they suggest that this speciation mode is uncommon in the genus and possibly in nature.

Evolution of intrinsic reproductive isolation among four North American populations of Rhagoletis pomonella (Diptera: Tephritidae)

Across its range in North America, four geographically separated, ecologically and genetically diverged populations of hawthorn (Crataegus)-infesting Rhagoletis pomonella (Diptera: Tephritidae) flies inhabit the Eje Volcanico Trans Mexicano (EVTM), the Sierra Madre Oriental (SMO), the Chiapas Highlands (CHIS) and the USA. Here, we tested whether these four populations are reproductively isolated by any intrinsic, nonhost-related, pre- or postmating barriers to gene flow. Crossing experiments suggested that a low level of host-independent prezygotic isolation may exist between hawthorn flies from EVTM and the three other populations, but only with respect to a slight reduction in copulation duration in EVTM matings. Some evidence for postmating isolation was found, again primarily involving EVTM crossed to SMO, CHIS and US flies. Certain crosses produced no (SMO male × EVTM female) or few (EVTM male × CHIS female; CHIS male × SMO female) F1 hybrid offspring. F2 crosses were generally fertile, except for US male × CHIS female matings. Inherent reproductive isolation therefore appears to be quantitative rather than absolute between populations, as the possibility for gene flow exists through at least some combinations of mating among EVTM, SMO, CHIS and US flies. Our results are consistent with a recently advanced hypothesis that episodic introgression from Mexico into the USA has played a role in providing genetic variation, facilitating sympatric host race formation and the adaptive radiation of the R. pomonella sibling species' complex in the USA. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100, 213–223.

Does strong selection promote host specialisation and ecological speciation in insect herbivores? Evidence from Neochlamisus leaf beetles

1. Outstanding issues in evolutionary ecology include: (1) the prevalence of host‐associated ecological specialisation among herbivorous insects, and (2) the speciation processes contributing to the tremendous species‐level diversity of herbivores.2. Neochlamisus leaf beetle species are generally monophagous, but cumulatively employ a disparate array of host plants. These patterns occur in microcosm within the putatively oligophagous Neochlamisus bebbianae, which uses specific host species within each of six plant genera representing five families. I refer to all beetle populations associated with any one of these hosts as a particular ‘host form’.3. This paper describes aspects of Neochlamisus biology and some experimental findings relevant to both host‐use evolution and host specialisation in Neochlamisus, and to ecological divergence, reproductive isolation, and genetic differentiation among N. bebbianae host forms. It thereby reviews a research programme focused on ecological speciation.4. With respect to these host forms, our findings include: (1) ubiquitous host specialisation, (2) constraints on ‘multitasking’ that select for host specialisation, (3) ubiquitous ecological divergence, (4) pre‐ and post‐mating reproductive barriers, (5) genetic and environmental contributions to ecological divergence and reproductive isolation, (6) positive statistical associations between ecological divergence and reproductive isolation, (7) strongly differentiated loci despite gene flow, reflecting divergent host‐related selection, and (8) possible host race formation. We additionally describe a unique example of intimate host adaptation in another species, Neochlamisus platani.5. The intimate and discrete nature of herbivore–host associations may generate unusually strong (host‐related) selection that could simultaneous explain pronounced rates of herbivore specialisation and speciation.

Differences in Preference and Performance of the Water Lily Leaf Beetle,Galerucella nymphaeaePopulations on Native and Introduced Aquatic Plants

Plant invasions represent ecological opportunities for herbivorous insects able to exploit novel host plants. The availability of new hosts and rapid adaptations may lead to host race formation and ultimately speciation. We studied potential host race formation in the water lily leaf beetle, Galerucella nymphaeae, in response to invasion by water chestnut, Trapa natans, in eastern North America. This leaf beetle is well suited for such studies because previous work showed that different herbivore populations follow different "evolutionary pathways" and specialize locally in response to differences in habitat preferences and host plant availability. We compared host preference and performance of G. nymphaeae offspring originating from T. natans and offspring of individuals originating from an ancestral host Nuphar lutea, yellow water lily, on T. natans and three native hosts (N. lutea, Nympheae odorata, and Brasenia schreberi). Regardless of origin (Trapa or Nuphar), adults strongly preferred their native host, N. lutea, over T. natans. Although laboratory survival rates (larva to pupa) were extremely high (80%) regardless of origin or host offered, survival rates in a common garden were greatly reduced, particularly for T. natans (24%) and to a lesser extent on N. lutea (54%), regardless of beetle origin. Larval drowning during more frequent leaf changes when developing on small Trapa leaves seems to be responsible for this difference. Preference of females for N. lutea is beneficial considering the much higher larval survival on the ancestral host. Abundant T. natans where the plant is invasive provides an alternative food source that beetles can use after egg/larval loads on their preferred host reach carrying capacity, but this utilization comes at a cost of high larval mortality.

Genetic Differentiation between Resistance Phenotypes in the Phytophagous Flea Beetle,Phyllotreta nemorum

The flea beetle Phyllotreta nemorum L. (Coleoptera: Chrysomelidae) is genetically polymorphic for resistance against the defences of one of its host plants, Barbarea vulgaris R.Br. (Brassicales: Brassicaceae). Whereas resistant flea beetles are able to use B. vulgaris as well as other cruciferous plants as food, non-resistant beetles cannot survive on B. vulgaris. This limitation to host plant use of non-resistant beetles could potentially lead to asymmetric gene flow and some degree of genetic isolation between the different resistance-genotypes. Therefore, we studied the extent of genetic differentiation at neutral allozyme loci between samples of flea beetles that were collected at different locations and first tested for resistance phenotype. Since earlier work has shown a weak, but significant, effect of geographical distance between the samples on their genetic differentiation, in the present study variation at the neutral allozyme loci in P. nemorum was partitioned between geographical distance and resistance-phenotype. Both sources independently contributed statistically significantly to population differentiation. Thus, there appears to be a limitation to genetic exchange between the resistant and non-resistant flea beetles when corrections are made for their geographic differentiation. This is consistent with the presence of some degree of host race formation in this flea beetle.

Recent evolution of host‐associated divergence in the seabird tick Ixodes uriae

Ecological interactions are an important source of rapid evolutionary change and thus may generate a significant portion of novel biodiversity. Such changes may be particularly prevalent in parasites, where hosts can induce strong selection for adaptation. To understand the relative frequency at which host-associated divergences occur, it is essential to examine the evolutionary history of the divergence process, particularly when it is occurring over large geographical scales where both geographical and host-associated isolation may playa part. In this study, we use population genetics and phylogeography to study the evolutionary history of host-associated divergence in the seabird tick Ixodes uriae (Acari, Ixodidae). We compare results from microsatellite markers that reflect more ecological timescales with a conserved mitochondrial gene (COIII) that reflects more ancient divergence events. Population structure based on microsatellites showed clear evidence of host-associated divergence in all colonies examined. However, isolated populations of the same host type did not always group together in overall analyses and the genetic differentiation among sympatric host races was highly variable. In contrast, little host or geographical structure was found for the mitochondrial gene fragment. These results suggest that host race formation in I. uriae is a recent phenomenon, that it may have occurred several times and that local interactions are at different points in the divergence process. Rapid divergence in I. uriae implies a strong interaction with its local host species, an interaction that will alter the ecological dynamics of the system and modify the epidemiological landscape of circulating micropathogens.

Phylogenetic analyses of the leaf beetle genus Galerucella: Evidence for host switching at speciation?

It is still the subject of lively debate whether sympatric speciation is a general mode of speciation as opposed to allopatric speciation. In herbivorous insects, host switching, i.e. colonization of, and adaptation to, a new host by a herbivore, has been proposed as one of the driving mechanisms of sympatric speciation. Evidence for host switching as a speciation driving mechanism can be inferred from phylogenies of herbivores and host plants: if the host plant phylogeny is randomly distributed over the herbivore phylogeny, this indicates host switching. The Chrysomelid beetle genus Galerucella is a good taxon to study for evidence of host switching, because several closely related Galerucella species form sympatric species complexes associated with various unrelated plant species. Here we present the phylogenetic relationships of 10 species in the genus Galerucella, based on the mitochondrial gene fragments of the NADH-2 (410 bp) and CO-I (659 bp) genes, and analyzed with Bayesian, Maximum Likelihood and Maximum Parsimony methods. The resulting molecular phylogenetic tree proved to be largely congruent with morphologically based taxonomy. The host-associated taxa of the Galerucella nymphaeae species complex are not defined as distinct gene pools under the phylogenetic species concept (PSC), however, the species complex as a whole is. Two results indicate the contribution of host switching to the speciation of Galerucella: (1) the host-associated taxa of the G. nymphaeae species complex have diverged very recently and (2) constrained ML analyses showed that host use constraints led to a significantly different Galerucella tree compared to unconstrained analyses. This evidence for host switching, together with the observation that several sister taxa using unrelated host plants live in sympatry, suggests that sympatric speciation by host race formation can be an important mode of speciation in this genus.

Host‐race formation: promoted by phenology, constrained by heritability

Host-race formation is promoted by genetic trade-offs in the ability of herbivores to use alternate hosts, including trade-offs due to differential timing of host-plant availability. We examined the role of phenology in limiting host-plant use in the goldenrod gall fly (Eurosta solidaginis) by determining: (1) whether phenology limits alternate host use, leading to a trade-off that could cause divergent selection on Eurosta emergence time and (2) whether Eurosta has the genetic capacity to respond to such selection in the face of existing environmental variation. Experiments demonstrated that oviposition and gall induction on the alternate host, Solidago canadensis, were the highest on young plants, whereas the highest levels of gall induction on the normal host, Solidago gigantea, occurred on intermediate-age plants. These findings indicate a phenological trade-off for host-plant use that sets up the possibility of divergent selection on emergence time. Heritability, estimated by parent-offspring regression, indicated that host-race formation is impeded by the amount of genetic variation, relative to environmental, for emergence time.

Infestation of a novel host plant by Tephritis conura (Diptera: Tephritidae) in northern Britain: host-range expansion or host shift?

The addition of a novel host plant to a phytophagous insect's diet may result in subsequent host-plant specialisation, and is believed to be a key cause for speciation in this trophic group. In northern Britain, the tephritid fly Tephritis conura has experienced a unique host-plant expansion, from the melancholy thistle Cirsium heterophyllum to the marsh thistle C. palustre. Here, we examine whether the incorporation of C. palustre in the repertoire of British T. conura flies has caused genetic divergence between populations infesting the old host and the novel host, and how British populations differ from populations infesting C. heterophyllum in continental Europe where C. palustre is not infested. No evidence for restricted gene flow among British C. palustre and C. heterophyllum flies was found. Significant differentiation between British and continental T. conura was found at only one allozyme locus, hexokinase, and caused by a new allele, Hex_95. Hexokinase is related to host-race formation in continental European flies infesting C. heterophyllum and C. oleraceum, and might be linked to loci determining host choice. Based on morphological and phenological data from previous studies, we suggest that T. conura in Britain has adapted to the novel host but that host-race formation is impeded by similar plant phenologies.