Cophylogenetic assessment of New World warblers (Parulidae) and their symbiotic feather mites (Proctophyllodidae)

Host–symbiont relationships are ubiquitous in nature, yet evolutionary and ecological processes that shape these intricate associations are often poorly understood. All orders of birds engage in symbioses with feather mites, which are ectosymbiotic arthropods that spend their entire life on hosts. Due to their permanent obligatory association with hosts, limited dispersal and primarily vertical transmission, we hypothesized that the cospeciation between feather mites and hosts within one avian family (Parulidae) would be perfect (strict cospeciation). We assessed cophylogenetic patterns and tested for congruence between species in two confamiliar feather mite genera (Proctophyllodidae: Proctophyllodes, Amerodectes) found on 13 species of migratory warblers (and one other closely related migratory species) in the eastern United States. Based on COI sequence data, we found three Proctophyllodes lineages and six Amerodectes lineages. Distance‐ and event‐based cophylogenetic analyses suggested different cophylogenetic trajectories of the two mite genera, and although some associations were significant, there was little overall evidence supporting strict cospeciation. Host switching is likely responsible for incongruent phylogenies. In one case, we documented prairie warblers Setophaga discolor harboring two mite species of the same genus. Most interestingly, we found strong evidence that host ecology may influence the likelihood of host switching occurring. For example, we documented relatively distantly related ground‐nesting hosts (ovenbird Seiurus aurocapilla and Kentucky warbler Geothlypis formosa) sharing a single mite species, while other birds are shrub/canopy or cavity nesters. Overall, our results suggest that cospeciation is not the case for feather mites and parulid hosts at this fine phylogenetic scale, and raise the question if cospeciation applies for other symbiotic systems involving hosts that have complex life histories. We also provide preliminary evidence that incorporating host ecological traits into cophylogenetic analyses may be useful for understanding how symbiotic systems have evolved.     

Habitat distribution of migratory and sedentary blackcaps Sylvia atricapilla wintering in southern Iberia: a morphological and biogeochemical approach

In migratory species, the way in which conspecifics from different breeding populations are distributed during the non‐breeding period is important from and ecological, evolutionary and conservation perspective, but such knowledge is still limited for most species. Migratory and sedentary blackcaps Sylvia atricapilla wintering in southern Spain can occupy two habitat types: forests and shrublands. According to earlier studies, blackcaps prefer forests over shrublands, and residents remain nearly restricted to forests. However, whether migrants with different breeding origin occupy the two habitats differently is unknown. We used morphological and biogeochemical data (hydrogen isotope ratios measured on feathers: δ²H_f), which show variation along the breeding range of the species, to answer this question. Isotope analyses supported the reliability of morphology as a method for distinguishing between migratory and sedentary blackcaps in sympatry, showing that sedentary individuals are rare in shrublands while migratory ones are abundant in both habitat types. However, migratory blackcaps scored similar δ²H_f values in forests and shrublands, and neither did vary in structural size or flight morphology between habitats. Our study suggests that migrants from a wide range of breeding origins end up mixing between forests and shrublands, which may explain the patterns of variation in space and time in the abundance of blackcaps in this area, and supports the view that inequalities may arise among migrants with the same origin but wintering in different habitats. Such inequalities might carry over into other stages of blackcaps’ life cycle contributing to the regulation of its migratory populations.     

The effect of water availability on fuel deposition of two staging Sylvia warblers

In order to succeed in crossing extensive ecological barriers, migratory birds usually deposit fuel en route. High rates of fuel deposition may enable birds to shorten their total migration time and are therefore advantageous for time‐minimizing migrants. Several studies have suggested that water provision may increase food utilization in non‐migratory birds. The goal of this study was to test the influence of water availability on the fuel deposition of en route migratory passerines. We studied fuel deposition of blackcaps Sylvia atricapilla and lesser whitethroats S. curruca staging in a plantation of Mount Atlas gum‐tree Pistacia atlantica in the northern Negev desert, Israel, during the autumns of 2000 and 2002. We manipulated water availability at the site and measured the effect of water supplementation on fuel deposition of birds of both species. We found that when water was available, blackcaps had higher fuel loads and higher fuel deposition rates than during control trials. However, water availability had no effect on fuel deposition of lesser whitethroats. Species‐specific differences in adaptations to arid conditions, reflected in the species’ winter habitat preferences, may be responsible for the between‐species dissimilarity in responding to water provision. We suggest that water availability may have strong ecological and evolutionary consequences for birds migrating through arid environments, by its possible effect on bird behavior and physiology.     

Contribution of oribatid and mesostigmatid soil mites in ecologically based estimates of global species richness

Erwin’s method for estimating total global species richness assumes some host‐specificity among the canopy arthropods. This study examined possible host habitat specialization in two major groups of soil arthropods, the oribatid and mesostigmatid mites, by sampling beneath three tree species: Eucalyptus pilularis Smith, Eucalyptus propinqua Deane and Maiden and Allocasuarina torulosa (Aiton) L. Johnson. The sample sites were in the Lansdowne State Forest, New South Wales, Australia and the three tree species were selected on the basis of their known differential effects on soil. Sampling was conducted over three seasons, and 79 oribatid and 34 mesostigmatid species were identified from 25 196 and 3634 individuals, respectively. Tree species had little effect on mite species composition with only three oribatid species and no mesostigmatid species identified as host‐habitat specialists using a niche breadth measure. Of mite species found under E. pilularis, E. propinqua and A. torulosa trees, 2%, 1% and 0% were defined as host‐habitat specialists, respectively. In contrast, tree species had significant and consistent effects on mite community structure, which differed in relative abundance of the oribatid species, their size class distributions and species rankings. In the mesostigmatid communities, there was a difference in the ranking of the mite species among tree species. Although it was demonstrated that tree species have an impact on the soil environment, the differences between tree species were insufficient to change species composition. The low degree of host‐habitat specialization suggested that other factors were more important for determining mite species composition at a site, and soil mite host‐habitat specialization may not make a large contribution to estimates of total global species richness using methods such as those proposed by Erwin (1982) .     

Variation in the mechanical properties of flight feathers of the blackcap Sylvia atricapilla in relation to migration

Migration causes temporal and energetic constraints during plumage development, which can compromise feather structure and function. In turn, given the importance of a good quality of flight feathers in migratory movements, selection may have favoured the synthesis of feathers with better mechanical properties than expected from a feather production constrained by migration necessities. However, no study has assessed whether migratory behaviour affects the relationship between the mechanical properties of feathers and their structural characteristics. We analysed bending stiffness (a feather mechanical property which is relevant to birds’ flight), rachis width and mass (two main determinants of variation in bending stiffness) of wing and tail feathers in migratory and sedentary blackcaps Sylvia atricapilla. Migratory blackcaps produced feathers with a narrower rachis in both wing and tail, but their feathers were not significantly lighter; in addition, bending stiffness was higher in migratory blackcaps than in sedentary blackcaps. Such unexpected result for bending stiffness remained when we statistically controlled for individual variation in rachis width and feather mass, which suggests the existence of specific mechanisms that help migratory blackcaps to improve the mechanical behaviour of their feathers under migration constraints.     

Historical diversification of migration patterns in a passerine bird

Migratory strategies of birds require complex orientation mechanisms, morphological adaptations, and life-history adjustments. From an evolutionary perspective, it is important to know how fast this complex combination of traits can evolve. We analyzed mitochondrial control-region DNA sequences in 241 blackcaps (Sylvia atricapilla) from 12 populations with different migratory behaviors. The sample included sedentary populations in Europe and Atlantic archipelagos and migratory populations with different distances of migration, from regional to intercontinental migrations, and different heading directions (due to a migratory divide in central Europe). There was no genetic structure between migratory and sedentary populations, or among populations from different biogeographic areas (Atlantic islands, the Iberian Peninsula, or the continent), however we found evidence of a genetic structure when comparing populations located on either side of the migratory divide. These findings support an independent evolution of highly divergent migratory strategies in blackcaps, occurring after a postglacial colonization of the continent along western and eastern routes. Accordingly, mismatch-distribution analyses suggested an expansion of blackcaps from a very small population size, and time estimates dated such an expansion during the last postglacial period. However, the populations in Gibraltar, located in a putative Mediterranean refuge, appeared to be independent of these processes, showing evidence of restricted gene flow with other populations and demonstrating insignificant historical changes in effective population size. Our results show that the interruption of gene flow between migratory and sedentary populations is not necessary for the maintenance of such a polymorphism, and that even the most divergent migratory strategies of a bird species are susceptible to evolution in response to historical environmental changes.     

Feather mite abundance varies but symbiotic nature of mite‐host relationship does not differ between two ecologically dissimilar warblers

Feather mites are obligatory ectosymbionts of birds that primarily feed on the oily secretions from the uropygial gland. Feather mite abundance varies within and among host species and has various effects on host condition and fitness, but there is little consensus on factors that drive variation of this symbiotic system. We tested hypotheses regarding how within‐species and among‐species traits explain variation in both (1) mite abundance and (2) relationships between mite abundance and host body condition and components of host fitness (reproductive performance and apparent annual survival). We focused on two closely related (Parulidae), but ecologically distinct, species: Setophaga cerulea (Cerulean Warbler), a canopy dwelling open‐cup nester, and Protonotaria citrea (Prothonotary Warbler), an understory dwelling, cavity nester. We predicted that feather mites would be more abundant on and have a more parasitic relationship with P. citrea, and within P. citrea, females and older individuals would harbor greater mite abundances. We captured, took body measurements, quantified feather mite abundance on individuals’ primaries and rectrices, and monitored individuals and their nests to estimate fitness. Feather mite abundance differed by species, but in the opposite direction of our prediction. There was no relationship between mite abundance and any measure of body condition or fitness for either species or sex (also contrary to our predictions). Our results suggest that species biology and ecological context may influence mite abundance on hosts. However, this pattern does not extend to differential effects of mites on measures of host body condition or fitness.     

Detecting ancient codispersals and host shifts by double dating of host and parasite phylogenies: Application in proctophyllodid feather mites associated with passerine birds

Inferring cophylogeographic events requires matching the timing of these events on both host and symbiont (e.g., parasites) phylogenies because divergences of hosts and their symbionts may not temporally coincide, and host switches may occur. We investigate a large radiation of birds (Passeriformes) and their permanent symbionts, the proctophyllodid feather mites (117 species from 116 bird species; six genes, 11,468 nt aligned) using two time‐calibration strategies for mites: fossils only and host phylogeography only. Out of 10 putative cophylogeographic events 4 agree in timing for both symbiont and host events being synchronous co‐origins or codispersals; three were based on host shifts, but agree in timing being very close to the origin of modern hosts; two disagree; and one large basal mite split was seemingly independent from host phylogeography. Among these events was an ancient (21–25.3 Mya), synchronous codispersal from the Old World leading to the origin and diversifications of New World emberizoid passerids and their mites, the thraupis + quadratus species groups of Proctophyllodes. Our framework offers a more robust detection of host and symbiont cophylogeographic events (as compared to host‐symbiont reconciliation analysis and using host phylogeography for time‐calibration) and provides independent data for testing alternative hypotheses on timing of host diversification and dispersal.     

Postglacial northward expansion and genetic differentiation between migratory and sedentary populations of the broad‐tailed hummingbird (Selasphorus platycercus)

Unlike other migratory hummingbirds in North America, the broad‐tailed hummingbird (Selasphorus platycercus) exhibits both long‐distance migratory behaviour in the USA and sedentary behaviour in Mexico and Guatemala. We examined the evolution of migration linked to its northward expansion using a multiperspective approach. We analysed variation in morphology, mitochondrial and nuclear DNA, estimated migration rates between migratory and sedentary populations, compared divergence times with the occurrence of Quaternary climate events and constructed species distribution models to predict where migratory and sedentary populations resided during the Last Glacial Maximum (LGM) and Last Interglacial (LIG) events. Our results are consistent with a recent northward population expansion driven by migration from southern sedentary populations. Phylogeographical analyses and population genetics methods revealed that migratory populations in the USA and sedentary populations in Mexico of the platycercus subspecies form one admixed population, and that sedentary populations from southern Mexico and Guatemala (guatemalae) undertook independent evolutionary trajectories. Species distribution modelling revealed that the species is a niche tracker and that the climate conditions associated with modern obligate migrants in the USA were not present during the LIG, which provides indirect evidence for recent migratory behaviour in broad‐tailed hummingbirds on the temporal scale of glacial cycles. The finding that platycercus hummingbirds form one genetic population and that suitable habitat for migratory populations was observed in eastern Mexico during the LIG also suggests that the conservation of overwintering sites is crucial for obligate migratory populations currently facing climate change effects.     

Identity,prevalence and intensity of infestation with wing feather mites on birds (Passeriformes) from the Setubal Peninsula of Portugal

The results of a 4 year project investigating species of mites infesting wing primary feathers on 21 species of Passeriformes are reported. The majority of species were identified as belonging to the genus Proctophyllodes Robin, 1877 with one new host record. In addition Pteronyssoides obscurus Berlese 1884 was found on European swallows, also a new host record. A novel method to enable quantification of mite intensities without causing harm to the birds was devised and evaluated. This relied on visual inspection of wing primary feathers and assignment of subjective infestation scores to individual feathers, the sum of the individual scores comprising the primary feather total mite infestation score (PTMIS). Comparisons between species revealed that birds could be grouped into four categoris depending on their infestation intensity with mites. Swallows, sand martins and greenfinches showed the highest prevalence and most intense infestations (mean PTMIS 6.8). Blackbirds, blackcaps, serins, goldfinches, Cetti's warblers, great tits and house sparrows showed moderate levels of infestation with prevalence in the range 60–90.9% but a mean PTMIS lower than in the former group (1.6–5.8). The third group comprised Sardinian warblers, nightingales and short-toed tree creepers and was characterized by a prevalence of mites 40% and a mean PTMIS of 0.6–1.4. The final group, representing wrens, chiffchaffs, fan-tailed warblers and waxbills were without detectable mites, the only exception being wrens on which mites were identified in only three birds of the 32 sampled. These results are interpreted in the light of published information and possible explanations for the observations are discussed.     

The evolution of dispersal polymorphisms in insects: The influence of habitats,host plants and mates

Wing-dimorphic, delphacid planthoppers were used to test hypotheses concerning the effects of habitat persistence and architectural complexity on the occurrence of dispersal. For reasons concerning both the durational stability of the habitat and the reduced availability of mates, selection has favored high levels of dispersal in species occupying temporary habitats. Flightlessness predominates in species occupying persistent habitats, and is promoted by a phenotypic trade-off between reproductive success and flight capability. Wings are retained in tree-inhabiting species, probably for reasons concerning the more effective negotiation of three-dimensional habitats. In contrast, flightlessness is characteristic of those species inhabiting low profile host plants. For several delphacid genera, migratory species are larger than their sedentary congeners. Because body size and fecundity are positively related in planthoppers, the large body size observed in migratory taxa may result from selection for increased fecundity in colonizing species.     

Habitat segregation by breeding origin in the declining populations of European Robins wintering in southern Iberia

Mediterranean woodlands and associated shrub formations of southern Iberia are key habitats for conservation of migratory birds. In some bird species, migratory and sedentary conspecifics meet in these areas during winter, but our understanding of how each population group is distributed over available habitats and the factors that determine their spatial organization are still unclear. This seriously limits our ability to assess their vulnerability to ongoing environmental changes affecting wintering habitats in this region. We used hydrogen isotopic signatures of feathers (δD_f) to shed light on the habitat distribution of seasonally sympatric European Robin Erithacus rubecula populations wintering in Campo de Gibraltar that are currently facing a drastic decline. In contrast to previous studies that used morphological methods to distinguish the migratory behaviour of wintering Robins in this area, our isotopic approach revealed that sedentary Robins were not outcompeted upon the arrival of migrants and remained in the woodlands where they reproduce, which agrees with results obtained in other regions. Interestingly, we also found that migratory Robins with a closer breeding origin (higher δD_f values) had a higher probability of occupying woodlands than did migrants coming from further away. Overall, our results suggest that the role of breeding origin in shaping the distribution of Robins during winter in Campo de Gibraltar is more relevant than the effects of sex, age or body size, which might facilitate the evolution of local adaptations for the exploitation of each habitat type.     

Migratory behaviour affects the trade-off between feather growth rate and feather quality in a passerine bird

Migratory birds have less time for moulting than sedentary birds, which may force them to produce their feathers faster at the expense of reducing feather quality. However, the effects of migration on the trade-off between moult speed and plumage quality remain to be studied in natural populations. We analysed the relationship between growth rate and quality of individual feathers, taking advantage of natural variation between migratory and sedentary populations of blackcaps Sylvia atricapilla . As predicted by life-history theory, individual blackcaps showed variable individual quality, which was revealed by positive correlations between feather growth rate and feather mass within populations. However, migrants grew up their feathers faster, producing lighter feathers than sedentary blackcaps. These results support the idea that feather growth rate and feather quality are traded against each other in blackcaps. Such a trade-off is apparently caused by different selection associated to migratory and sedentary life styles, which opens new insights into the diversification of moult patterns in birds.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 98–105.     

MULTITRAIT,HOST‐ASSOCIATED DIVERGENCE AMONG SETS OF BUTTERFLY POPULATIONS: IMPLICATIONS FOR REPRODUCTIVE ISOLATION AND ECOLOGICAL SPECIATION

When populations use different resources, they tend to diverge in traits that affect performance on those resources. The extent and complexity of divergence that is achieved will depend on gene flow, genetic constraints, and the character of divergent selection. We describe divergent host adaptation among Californian populations of the Melitaeine butterfly, Euphydryas editha. Divergence in seven traits created parallel phenotypic suites, each suite associated with the use of a different host species, either Collinsia torreyi or Pedicularis semibarbata. The suites involved alighting responses of adults (probably to visual stimuli), chemosensory responses to leaf surfaces, vertical positioning of adults and larvae (probably due to geotaxis), partitioning of reproductive effort among clutches, and larval performance. Remarkably, the divergent suites did not occur sympatrically, despite ubiquitous co‐occurrence of the hosts, and we know of only one site where any Collinsia species is used sympatrically with another host. In contrast, E. editha often uses two host genera sympatrically when neither of them is Collinsia. We suggest that adaptation to Collinsia is incompatible with adaptation to other hosts and may generate extrinsic postzygotic reproductive isolation among populations. Despite the apparent rarity of host‐shift‐associated speciation in Melitaeine butterflies, adoption of Collinsia as a host may lead to allopatric ecological speciation.     

Contrasting effects of Wolbachia on cytoplasmic incompatibility and fecundity in the haplodiploid mite Tetranychus urticae

Recent studies on Wolbachia‐induced incompatibility in haplodiploid insects and mites have revealed a diversity of cytoplasmic incompatibility (CI) patterns among host species. Here, we report intraspecific diversity in CI expression among four strains of the arrhenotokous mite Tetranychus urticae and in T. turkestani. Variability of CI expression within T. urticae ranged from no CI to complete CI, and included either female embryonic mortality or male conversion types of CI. A fecundity cost attributed to the infection with the high‐CI Wolbachia strain was the highest ever recorded for Wolbachia (?80 to ?100% decrease). Sequence polymorphism at a 550‐bp‐portion of Wolbachia wsp gene revealed two clusters distant by 21%, one of which included three Wolbachia strains infecting mite populations sampled from the same host‐plant species, but showing distinct CI patterns. These data are discussed in the light of theoretical predictions on the evolutionary pathways followed in this symbiotic interaction.     

Individual differences in migratory behavior shape population genetic structure and microhabitat choice in sympatric blackcaps (Sylvia atricapilla)

In migratory birds, traits such as orientation and distance are known to have a strong genetic background, and they often exhibit considerable within‐population variation. How this variation relates to evolutionary responses to ongoing selection is unknown because the underlying mechanisms that translate environmental changes into population genetic changes are unclear. We show that within‐population genetic structure in southern German blackcaps (Sylvia atricapilla) is related to individual differences in migratory behavior. Our 3‐year study revealed a positive correlation between individual migratory origins, denoted via isotope (δ²H) values, and genetic distances. Genetic diversity and admixture differed not only across a recently established migratory polymorphism with NW‐ and SW‐migrating birds but also across δ²H clusters within the same migratory route. Our results suggest assortment based on individual migratory origins which would facilitate evolutionary responses. We scrutinized arrival times and microhabitat choice as potential mechanisms mediating between individual variation in migratory behavior and assortment. We found significant support that microhabitat choice, rather than timing of arrival, is associated with individual variation in migratory origins. Moreover, examining genetic diversity across the migratory divide, we found migrants following the NW route to be genetically more distinct from each other compared with migrants following the traditional SW route. Our study suggests that migratory behavior shapes population genetic structure in blackcaps not only across the migratory divide but also on an individual level independent of the divide. Thus, within‐population variation in migratory behavior might play an important role in translating environmental change into genetic change.     

Effects of habitat heterogeneity and local adaptation on the body condition of a forest passerine at the edge of its distributional range

We studied fluctuating asymmetry and feather growth rates as indicators of fitness of blackcaps ( Sylvia atricapilla ) breeding at the border of their distribution range in the Iberian Peninsula. Iberian blackcaps increase their abundance with increasing rainfall and ground cover of brambles. In border habitats, they are sedentary and morphologically different from migrants, suggesting that they could be adapted to peripheral conditions. We tested whether juvenile body condition depends on (1) distance from the centre of the range, (2) mean precipitation or (3) bramble cover. Controlling for environmental variation, we tested for differences between migratory and sedentary populations. Body condition varied across the Iberian gradient in parallel with changes in precipitation. Controlling for this effect, sedentary populations were in better body condition than migratory populations. Our results support the idea that environmental heterogeneity causes fitness to fluctuate across species' ranges, and also that local adaptation may mean that peripheral populations are more than a 'tail end' of the species.  © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 78 , 479–488.     

Symbiotic feather mites synchronize dispersal and population growth with host sociality and migratory disposition

Some symbiotic taxa may have evolved to track changes in the level and quality of food resources provided by the host to increase reproduction and dispersal. As a consequence, some ectosymbionts synchronize their reproduction and activity with particular stages of their host's living cycle. In this article we examined temporal patterns of variation in prevalence and abundance of feather mites living on pre‐migratory barn swallows Hirundo rustica. Feather mites in the lineages Pterolichoidea and Analgoidea are the most common arthropod ectosymbionts living at the expenses of feather oil. We investigated whether the seasonal variations in levels of several measures of physiological condition associated with host migration were related to changes in prevalence and abundance of mites. The results suggest that the variation in prevalence of feather mites, and thus probably the mode of acquisition and dispersal of these symbionts, is linked to an increase in host sociality before migration. Physiological dynamics of hosts after the breeding season point at two clearly identifiable periods: a post‐breeding period when physiological condition remains stationary or decreases, and a pre‐migratory period characterized by a rapid increase in several measures of physiological condition. Mite population dynamics were synchronized with migratory disposition during the period of highest host gregariousness. These synchronized processes occurred in both study years, although dynamics of migratory disposition and mite prevalence and abundance differ somewhat between years for adult and juvenile hosts. Mite population increase before host migration may be a response to a higher quantity of food provided by the host, namely oil from the urpoygial gland which is stimulated by hormones. Therefore, mites might have evolved to adjust their reproduction to the time when they have more chance of dispersal through horizontal transmission. In addition, body mass of juvenile and adult hosts were positively related with mite abundance in both years after allowing for several influencing factors. Body mass variation may reflect adequately fitness of host or their current physiological state, for instance, differences in the secretion of lipids on feathers or a more adequate microclimate to these symbionts.     

The wheat curl mite Aceria tosichella (Acari: Eriophyoidea) is a complex of cryptic lineages with divergent host ranges: evidence from molecular and plant bioassay data

Aceria tosichella (the wheat curl mite, WCM) is a global pest of wheat and other cereals, causing losses by direct damage, as well as the transmission of plant viruses. The mite is considered to have an unusually wide host range for an eriophyoid species. The present study tested the commonly held assumption that WCM is a single, highly polyphagous species by assessing the host range of genetically distinct lineages of WCM occurring in Poland on different host plants. Genotyping was performed by analyzing nucleotide sequence data from fragments of the mitochondrial cytochrome c oxidase subunit I (COI) and the nuclear D2 region of 28S rDNA. Mean between‐lineage distance estimated using COI data was found to be one order of magnitude greater than the within‐clade lineage and, in some cases, comparable to distances between WCM lineages and a congeneric outgroup species. Host acceptance was tested by quantifying population growth for different WCM mitochondrial (mt)DNA lineages when transferred from source host plants to test plants. These experiments revealed significant differences in host colonization ability between mtDNA lineages, ranging from highly polyphagous to more host‐specific. The present study reveals that WCM is composed of several discrete genetic lineages with divergent host‐acceptance and specificity traits. Genetic variation for host acceptance within A. tosichella s.l. may act as a reproductive barrier between these lineages, most of which had narrow host ranges. Two lineages appear to have high pest potential on cereals, whereas several others appear to specialize on wild grass species. We conclude that WCM is not a homogeneous species comprising polyphagous panmictic populations rather it is a complex of genetically distinct lineages with variable host ranges and therefore variable pest potential. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 165–180.