Allopatric Host Research Articles

Local adaptation of a generalist hemiparasitic plant to one of its potential host plants

Coevolution is often found in parasite–host interactions, but has not yet been described for hemiparasitic plants and their hosts. Root hemiparasites like Rhinanthus alectorolophus perform photosynthesis but also parasitize other plant species, some of which (e.g. Plantago lanceolata) may defend themselves against parasite attack by blocking the haustoria of the parasites. We grew seedlings of the hemiparasite R. alectorolophus and the potential host P. lanceolata from seven grassland sites in a factorial design. To detect differences in host defence, we also included hosts from two ‘naïve' populations from regions where the parasite does not occur. R. alectorolophus grew consistently larger and had higher fitness with sympatric than with allopatric hosts, suggesting parasite adaptation to local host populations. Moreover, R. alectorolophus remained smallest with allopatric hosts from the same region and reached intermediate sizes with allopatric hosts from other regions or naïve hosts, suggesting host adaptation to parasites at the regional scale. Parasite presence reduced the size of the host plants already after four weeks, but only that of hosts with ‘experience' of the parasite, suggesting an early host response. Follow‐up experiments confirmed that parasites attach to hosts already after four weeks and hosts respond by changing belowground allocation patterns. However, parasite roots did not preferentially grow towards sympatric hosts. Our results suggest that local adaptation to hosts can occur even in generalist parasites and does not require specialization on individual hosts. We discuss the role of potential mechanisms, including variation in chemical signalling (early) and in host defence (late effects).

Do biological control agents adapt to local pest genotypes? A multiyear test across geographic scales.

Parasite local adaptation has been a major focus of (co)evolutionary research on host-parasite interactions. Studies of wild host-parasite systems frequently find that parasites paired with local, sympatric host genotypes perform better than parasites paired with allopatric host genotypes. In contrast, there are few such tests in biological control systems to establish whether biological control parasites commonly perform better on sympatric pest genotypes. This knowledge gap prevents the optimal design of biological control programs: strong local adaptation could argue for the use of sympatric parasites to achieve consistent pest control. To address this gap, we tested for local adaptation of the biological control bacterium Pasteuria penetrans to the root-knot nematode Meloidogyne arenaria, a global threat to a wide range of crops. We measured the probability and intensity of P. penetrans infection on sympatric and allopatric M. arenaria over the course of 4 years. Our design accounted for variation in adaptation across scales by conducting tests within and across fields, and we isolated the signature of parasite adaptation by comparing parasites collected over the course of the growing season. Our results are largely inconsistent with local adaptation of P. penetrans to M. arenaria: in 3 of 4 years, parasites performed similarly well in sympatric and allopatric combinations. In 1 year, however, infection probability was 28% higher for parasites paired with hosts from their sympatric plot, relative to parasites paired with hosts from other plots within the same field. These mixed results argue for population genetic data to characterize the scale of gene flow and genetic divergence in this system. Overall, our findings do not provide strong support for using P. penetrans from local fields to enhance biological control of Meloidogyne.

The role of adaptive resistance in a widespread freshwater mussel species

Abstract Organism features, such as size, weight, sex and age, among others, can influence the success of parasitism. For species that depend on a host to complete their life cycle, such as freshwater mussels (Bivalvia, Unionida), understanding how these traits influence parasitism can help guide management actions aimed at improving their reproductive success, ultimately benefitting their conservation. In this study, the physiological compatibility between the duck mussel Anodonta anatina and its host fish was investigated in a simultaneous infestation experiment on four (three native and one non‐native) fish species using sympatric and allopatric host strains. The success of glochidia metamorphosis in juvenile mussels was evaluated as a function of host species, strain and fish length. Successful metamorphosis was achieved in almost all fish species tested and differences were detected between allopatric and sympatric strains, with higher values found in allopatric hosts. Allopatric strains of primary hosts produce more juveniles than marginal hosts and non‐native species. In addition, larger fish have lower rates of metamorphosis. The results confirm recent findings showing high variation in the metamorphosis success of A. anatina glochidia among different native and non‐native hosts, both in the natural environment and in artificial laboratory infestations. Allopatric hosts were more suitable for encystment, demonstrating potential adaptive immunity, not yet observed in A. anatina. Nevertheless, A. anatina may temporarily exploit the naivety of smaller fish regardless of strain. Overall, the results suggest that mussel propagation, reproduction and reintroduction efforts should consider the mechanisms that influence adaptive immunity in fish, including differences in compatibility within and between populations of sympatric and allopatric host strains. This situation should be taken into account, as successful metamorphosis is an important factor in the selection of hosts for juvenile development, especially in efforts to propagate endangered mussel species.

Genetic diversification of an invasive honey bee ectoparasite across sympatric and allopatric host populations.

Invasive parasites are major threats to biodiversity. The honey bee ectoparasite, Varroa destructor, has shifted host and spread almost globally several decades ago. This pest is generally considered to be the main global threat to Western honey bees, Apis mellifera, although the damages it causes are not equivalent in all its new host's populations. Due to the high virulence of this parasite and the viruses it vectors, beekeepers generally rely on acaricide treatments to keep their colonies alive. However, some populations of A. mellifera can survive without anthropogenic mite control, through the expression of diverse resistance and tolerance traits. Such surviving colonies are currently found throughout the globe, with the biggest populations being found in Sub-Saharan Africa and Latin America. Recently, genetic differences between mite populations infesting surviving and treated A. mellifera colonies in Europe were found, suggesting that adaptations of honey bees drive mite evolution. Yet, the prevalence of such co-evolutionary adaptations in other invasive populations of V. destructor remain unknown. Using the previous data from Europe and novel genetic data from V. destructor populations in South America and Africa, we here investigated whether mites display signs of adaptations to different host populations of diverse origins and undergoing differing management. Our results show that, contrary to the differences previously documented in Europe, mites infesting treated and untreated honey bee populations in Africa and South America are genetically similar. However, strong levels of genetic differentiation were found when comparing mites across continents, suggesting ongoing allopatric speciation despite a recent spread from genetically homogenous lineages. This study provides novel insights into the co-evolution of V. destructor and A. mellifera, and confirms that these species are ideal to investigate coevolution in newly established host-parasite systems.

Fitness consequences of host colonization in two generalist fleas: Context-dependency and the effect of spatial co-occurrence.

We studied the fitness consequences of colonizing a novel host by experimental lines of fleas (Synosternus cleopatrae and Xenopsylla ramesis) maintained for 18-22 generations on the principal or novel (sympatric or allopatric) hosts via number, developmental success and size of the offspring of the fleas exploiting these hosts. We asked whether (a) fitness on non-principal hosts increases after prolonged maintenance; (b) the colonization success depends on the spatial co-occurrence of a flea and a host and (c) colonization of a novel host is accompanied by a decreased ability to exploit an original host. The ability of fleas to colonize novel hosts differed between species, with S. cleopatrae, but not X. ramesis, increasing its offspring production on novel hosts. Spatial co-occurrence did not affect colonization success. Maintenance on an alternative host was not accompanied by decreased adaptation to the original host. When fleas returned to the original host, their reproductive output was higher than that of their ancestors. We conclude that the success of colonizing a novel host is (a) context-dependent and varies between flea and host species and (b) not accompanied by the loss of ability to exploit an ancestral host but may lead to an increase in this ability.

Colonization of a novel host by fleas: changes in egg production and egg size

We studied the success of fleas, Synosternus cleopatrae and Xenopsylla ramesis, in switching to a novel host by establishing experimental lines maintained on different hosts for 18 generations. Fleas fed on principal (P-line) or novel hosts, either sympatric with (S-line) or allopatric to (A-line) a flea and its principal host, then we assessed their reproductive performance via the number and size of eggs. We compared reproductive performance between hosts within a line and between lines within a host asking: (a) whether fleas adapt to a novel host species after multiple generations; (b) if yes, whether the pattern of adaptation differs between novel host species sympatric with or allopatric to a flea and its principal host; and (c) adaptation to a novel host is accompanied with a loss of success in exploitation of an original host. Fleas from the S- and A-lines increased their egg production on a novel host (except X. ramesis from the S-line). S. cleopatrae from the S-line but not the A-line increased egg size on a novel host, whereas X. ramesis from the A-line but not the S-line produced larger eggs from a novel host. We found no indication of a loss of reproductive performance on the original host while adapting to a novel host. We conclude that fleas are able to switch rapidly to a new host with the pattern of a switch to either sympatric or an allopatric host depending on the identities of both flea and host species.

Patterns of Sphaeridiotrema pseudoglobulus infection in sympatric and allopatric hosts (Bithynia tentaculata) originating from widely separated sites across the USA.

In circumstances where populations of invasive species occur across variable landscapes, interactions among invaders, their parasites, and the surrounding environment may establish local coevolutionary trajectories for the participants. This can generate variable infection patterns when parasites interact with sympatric versus allopatric hosts. Identifying the potential for such patterns within an invasive-species framework is important for better predicting local infection outcomes and their subsequent impacts on the surrounding native community. To begin addressing this question, we exposed an invasive snail (Bithynia tentaculata) from two widely separated sites across the USA (Wisconsin and Montana) to the digenean parasite, Sphaeridiotrema pseudoglobulus, collected from Wisconsin. Parasite exposures generated high infection prevalences in both sympatric and allopatric snails. Furthermore, host survival, host growth, the proportion of patent snails, and the timing of patency did not differ between sympatric and allopatric combinations. Moreover, passaging parasites through snails of different origins had no effect on transmission success to subsequent hosts in the life cycle. However, the number of parasites emerging from snails and the pattern of their release varied based on snail origin. These latter observations suggest the potential for local adaptation in this system, but subsequent research is required to further substantiate this as a key factor underlying infection patterns in the association between S. pseudoglobulus and B. tentaculata.

The Role of Sympatric and Allopatric Speciation in the Origin of Biodiversity of Herbivorous Insects, with Palaearctic Species of the Genus Macropsis Lewis, 1836 Taken as an Example (Homoptera, Auchenorrhyncha, Cicadellidae, Eurymelinae, Macropsini)

Using 60 Palaearctic species of the leafhopper genus Macropsis as an example, the contribution of sympatric and allopatric speciation to the origin of herbivorous insect biodiversity was studied. According to the previously reconstructed evolutionary scenarios, in the temperate zone of the Palaearctic Region the genus Macropsis probably dispersed from east to west and formed 10 natural groups, with species within each group being similar both in morphological traits and in host specialization. The hypothetical speciation modes for each group can be largely inferred from geography. If related species replace each other in different zoogeographic regions, they are likely to have arisen as the result of allopatric speciation. If the species show overlapping distributions and live on the same host plant, they must have originated in allopatry and become secondarily sympatric due to subsequent dispersal. Finally, if related species have strongly overlapping ranges and populate similar habitats but differ in host specializations, this may indicate their origin as the result of sympatric speciation due to host plant shifts. For 31 species (51.7%), allopatric speciation seems to be the most likely or even the only possible scenario, whereas for 20 species (33.3%), the hypothesis of sympatric speciation is either convincingly substantiated or at least quite feasible. The origins of the remaining 9 species (15%) remain unknown. Therefore, sympatric speciation is not a rare phenomenon in Macropsis, but it certainly does not predominate over the allopatric mode. Thus, even within the same genus of herbivorous insects, speciation can occur in different ways: in the allopatric mode, when a host plant shift is possible but not essential, and in the sympatric mode, when divergence is triggered by the host shift. Cases of intraspecific differentiation, which can be considered as initial stages of speciation, are also discussed. These are allopatric color forms associated with the same host plant, allopatric color forms associated with different plants and, finally, forms living on different sympatric or allopatric hosts but showing no difference in coloration. It is shown that divergence is not always caused by transition to a new host plant even in specialized herbivores.

Quantitative trait loci involved in the reproductive success of a parasitoid wasp.

Dissecting the genetic basis of intraspecific variations in life history traits is essential to understand their evolution, notably for potential biocontrol agents. Such variations are observed in the endoparasitoid Cotesia typhae (Hymenoptera: Braconidae), specialized on the pest Sesamia nonagrioides (Lepidoptera: Noctuidae). Previously, we identified two strains of C. typhae that differed significantly for life history traits on an allopatric host population. To investigate the genetic basis underlying these phenotypic differences, we used a quantitative trait locus (QTL) approach based on restriction site-associated DNA markers. The characteristic of C. typhae reproduction allowed us generating sisters sharing almost the same genetic content, named clonal sibship. Crosses between individuals from the two strains were performed to generate F2 and F8 recombinant CSS. The genotypes of 181 clonal sibships were determined as well as the phenotypes of the corresponding 4,000 females. Informative markers were then used to build a high-quality genetic map. These 465 markers spanned a total length of 1,300cM and were organized in 10 linkage groups which corresponded to the number of C. typhae chromosomes. Three QTLs were detected for parasitism success and two for offspring number, while none were identified for sex ratio. The QTLs explained, respectively, 27.7% and 24.5% of the phenotypic variation observed. The gene content of the genomic intervals was investigated based on the genome of C. congregata and revealed 67 interesting candidates, as potentially involved in the studied traits, including components of the venom and of the symbiotic virus (bracovirus) shown to be necessary for parasitism success in related wasps.

Host-Dependent Differences in Replication Strategy of the Sulfolobus Spindle-Shaped Virus Strain SSV9 (a.k.a., SSVK1): Infection Profiles in Hosts of the Family Sulfolobaceae.

The Sulfolobus Spindle-shaped Virus (SSV) system has become a model for studying thermophilic virus biology, including archaeal host-virus interactions and biogeography. Several factors make the SSV system amenable to studying archaeal genetic mechanisms (e.g., CRISPRs) as well as virus-host interactions in high temperature acidic environments. Previously, we reported that SSVs exhibited differential infectivity on allopatric vs. sympatric hosts. We also noticed a wide host range for virus strain SSV9 (a.k.a., SSVK1). For decades, SSVs have been described as “non-lytic” double-stranded DNA viruses that infect species of the genus Sulfolobus and release virions via budding rather than host lysis. In this study, we show that SSVs infect hosts representing more than one genus of the family Sulfolobaceae in spot-on-lawn “halo” assays and in liquid culture infection assays. Growth curve analyses support the hypothesis that SSV9 virion release causes cell lysis. While SSV9 appears to lyse allopatric hosts, on a single sympatric host, SSV9 exhibits canonical non-lytic viral release historically reported SSVs. Therefore, the nature of SSV9 lytic-like behavior may be driven by allopatric evolution. The SSV9-infected host growth profile does not appear to be driven by multiplicity of infection (MOI). Greater stability of SSV9 vs. other SSVs (i.e., SSV1) in high temperature, low pH environments may contribute to higher transmission rates. However, neither higher transmission rate nor relative virulence in SSV9 infection seems to alter replication profile in susceptible hosts. Although it is known that CRISPR-Cas systems offer protection against viral infection in prokaryotes, CRISPRS are not reported to be a determinant of virus replication strategy. The mechanisms underlying SSV9 lytic-like behavior remain unknown and are the subject of ongoing investigations. These results suggest that genetic elements, potentially resulting from allopatric evolution, mediate distinct virus-host growth profiles of specific SSV-host strain pairings.

Host-dependent differences in replication and virion release strategy of the Sulfolobus Spindle-shaped Virus strain SSV9 (a.k.a., SSVK1): Lytic replication in hosts of the family Sulfolobaceae

The Sulfolobus Spindle-shaped Virus (SSV) system is a model for studying thermophilic archaeal virus biology. Several factors make the SSV system amenable to studying archaeal genetics and virus-host interactions in extreme environments. It has been shown that populations of Sulfolobus, the natural host, exhibit biogeographic structure. The acidic (pH<4.5) high temperature (65-88°C) habitats have low biodiversity, which diminishes prospects for host switch. SSVs and their hosts are readily cultured in liquid media and on plates. Given the wide geographic separation between various SSV-Sulfolobus habitats, the system is also amenable to studying allopatric versus sympatric virus-host interactions. We previously reported that SSVs exhibit differential infectivity on allopatric and sympatric hosts. We discovered a strikingly broad host-range for strain SSV9 (a.k.a., SSVK1). For decades, SSVs have been described as “non-lytic” dsDNA viruses that infect species of Sulfolobus and release virus particles via blebbing as a preferred strategy over host lysis (in reported laboratory infections). Here, we show, that SSVs infect more than one genus of the family Sulfolobaceae and, in allopatric hosts, SSV9 does not release virions via blebbing. Instead, SSV9 appears to lyse all susceptible allopatric hosts, while exhibiting canonical non-lytic virion release (historically reported for SSVs) on a single sympatric host. Lytic versus non-lytic virus release does not appear to be driven by multiplicity of infection. Data suggest that SSV9 is more stable than other SSVs in suspension; however, genetic substrates (e.g., CRISPR profiles) underlying non-lytic versus lytic virion release remain unresolved and are the subject of ongoing investigation.

Host plant adaptation during contemporary range expansion in the monarch butterfly.

Herbivores that have recently expanded their host plant ranges provide opportunities to test hypotheses about the evolution of host plant specialization. Here, we take advantage of the contemporary global range expansion of the monarch butterfly (Danaus plexippus) and conduct a reciprocal rearing experiment involving monarch populations with divergent host plant assemblages. Specifically, we ask the following questions: (1) Do geographically disparate populations of monarch butterflies show evidence for local adaptation to their host plants? If so, what processes contribute to this pattern? (2) How is dietary breadth related to performance across multiple host species in monarch populations? (3) Does the coefficient of variation in performance vary across sympatric versus allopatric hosts? We find evidence for local adaptation in larval growth rate and survival based on sympatric/allopatric contrasts. Migratory North American monarchs, which have comparatively broad host breadth, have higher mean performance than derived nonmigratory populations across all host plant species. Monarchs reared on their sympatric host plants show lower coefficient of variation in performance than monarchs reared on allopatric hosts. We focus our discussion on possible mechanisms contributing to local adaptation to novel host plants and potential explanations for the reduction in performance that we observed in derived monarch populations.

Climatic synchrony and increased outbreaks in allopatric populations of an invasive defoliator

Larch casebearer is an invasive defoliator in North America distributed within the regions of two allopatric hosts, eastern larch and western larch. Despite the establishment of a successful importation biological control program and ongoing parasitism by both native and introduced parasitoids, larch casebearer has recently undergone outbreaks on eastern larch and western larch. We analyzed defoliation data from aerial surveys to quantify spatiotemporal synchrony and found that defoliation by larch casebearer was synchronous both within and between eastern and western larch forests. We also analyzed monthly minimum temperatures across the study region and found that warming spring temperatures in March and cooling fall/winter temperatures in October through December were positively correlated at distances comparable to those between the allopatric outbreaks of larch casebearer. For allopatric populations with positively correlated population dynamics, climate is the most likely driver of synchrony. Thus, we suggest that a changing climate has facilitated recent outbreaks of larch casebearer.

Manipulating host resistance structure reveals impact of pathogen dispersal and environmental heterogeneity on epidemics.

Understanding how variation in hosts, parasites, and the environment shapes patterns of disease is key to predicting ecological and evolutionary outcomes of epidemics. Yet in spatially structured populations, variation in host resistance may be spatially confounded with variation in parasite dispersal and environmental factors that affect disease processes. To tease apart these disease drivers, we paired surveys of natural epidemics with experiments manipulating spatial variation in host susceptibility to infection. We mapped epidemics of the wind-dispersed powdery mildew pathogen Podosphaera plantaginis in five populations of its plant host, Plantago lanceolata. At 15 replicate sites within each population, we deployed groups of healthy potted 'sentinel' plants from five allopatric host lines. By tracking which sentinels became infected in the field and measuring pathogen connectivity and microclimate at those sites, we could test how variation in these factors affected disease when spatial variation in host resistance and soil conditions was minimized. We found that the prevalence and severity of sentinel infection varied over small spatial scales in the field populations, largely due to heterogeneity in pathogen prevalence on wild plants and unmeasured environmental factors. Microclimate was critical for disease spread only at the onset of epidemics, where humidity increased infection risk. Sentinels were more likely to become infected than initially healthy wild plants at a given field site. However, in a follow-up laboratory inoculation study we detected no significant differences between wild and sentinel plant lines in their qualitative susceptibility to pathogen isolates from the field populations, suggesting that primarily non-genetic differences between sentinel and wild hosts drove their differential infection rates in the field. Our study leverages a multi-faceted experimental approach to disentangle important biotic and abiotic drivers of disease patterns within wild populations.

Origin and macroevolution of micro-moths on sunken Hawaiian Islands.

The origins and evolution of Hawaiian biodiversity are a matter of controversy, and the mechanisms of lineage diversification for many organisms on this remote archipelago remain unclear. Here we focus on the poorly known endemic leaf-mining moth genus Philodoria (Lepidoptera, Gracillariidae), whose species feed on a diversity of Hawaiian plant lineages, many of which are critically endangered. We use anchored hybrid enrichment to assemble the first phylogenomic dataset (507 loci) for any Hawaiian animal taxon. To uncover the timing and pattern of diversification of these moths, we apply two frequently used dating calibration strategies, biogeographic calibrations and secondary calibrations. Island calibrations on their own resulted in much younger and unrealistic dates compared to strategies that relied on secondary calibrations. Philodoria probably originated on the now partially sunken islands of Laysan or Lisianski, approximately 21 Ma, and were associated with host plants in the families Ebenaceae, Malvaceae or Primulaceae. Major feeding groups associated with specific host-plant families originated soon after the plants colonized the islands. Allopatric isolation and host shifts, in concert and independently, probably play major roles in the diversification of Philodoria Our dating results indicate that Philodoria is among the oldest known Hawaiian arthropod lineages, and that island calibrations alone can lead to unrealistically young dates.

Epidemiological trade-off between intra- and interannual scales in the evolution of aggressiveness in a local plant pathogen population.

The efficiency of plant resistance to fungal pathogen populations is expected to decrease over time, due to their evolution with an increase in the frequency of virulent or highly aggressive strains. This dynamics may differ depending on the scale investigated (annual or pluriannual), particularly for annual crop pathogens with both sexual and asexual reproduction cycles. We assessed this time‐scale effect, by comparing aggressiveness changes in a local Zymoseptoria tritici population over an 8‐month cropping season and a 6‐year period of wheat monoculture. We collected two pairs of subpopulations to represent the annual and pluriannual scales: from leaf lesions at the beginning and end of a single annual epidemic and from crop debris at the beginning and end of a 6‐year period. We assessed two aggressiveness traits—latent period and lesion size—on sympatric and allopatric host varieties. A trend toward decreased latent period concomitant with a significant loss of variability was established during the course of the annual epidemic, but not over the 6‐year period. Furthermore, a significant cultivar effect (sympatric vs. allopatric) on the average aggressiveness of the isolates revealed host adaptation, arguing that the observed patterns could result from selection. We thus provide an experimental body of evidence of an epidemiological trade‐off between the intra‐ and interannual scales in the evolution of aggressiveness in a local plant pathogen population. More aggressive isolates were collected from upper leaves, on which disease severity is usually lower than on the lower part of the plants left in the field as crop debris after harvest. We suggest that these isolates play little role in sexual reproduction, due to an Allee effect (difficulty finding mates at low pathogen densities), particularly as the upper parts of the plant are removed from the field, explaining the lack of transmission of increases in aggressiveness between epidemics.

Habitat Heterogeneity, Host Population Structure, and Parasite Local Adaptation.

Reciprocal-transplant experiments have proven to be a powerful tool for detecting local adaptation (LA). More recently, reciprocal cross-inoculation experiments have been used to evaluate adaptation by parasites to their local host populations. These experiments are conceptually similar to reciprocal-transplant experiments, except that the "environment" (the host population) may have evolved in response to changes in the parasite population. Here, I use analytical tools and computer simulations to determine when parasites would be expected to be more infective to their local host populations than to allopatric host populations. The models assume that parasites have to genetically "match" their hosts in order to infect. I also assumed that different host clones were favored in different populations. When parasite virulence was low, clonal selection outweighed parasite-mediated selection, leading to low host diversity within populations and strong LA by parasites. At intermediate levels of virulence, parasite-mediated selection maintained high levels of host diversity within populations, which reduced or eliminated parasite LA. The loss of parasite LA was not associated with increased infectivity by parasites on allopatric hosts. Instead, the loss of LA was due to a reduction in infectivity of parasites on sympatric hosts. Finally, at high levels of parasite virulence, parasite-mediated selection led to oscillatory host dynamics and weak local adaption by parasites. Across all levels of virulence, the strength of parasite LA closely tracked the degree of host population structure (GST).

Relationship between oviposition, virulence gene expression and parasitism success in Cotesia typhae nov. sp. parasitoid strains.

Studying mechanisms that drive host adaptation in parasitoids is crucial for the efficient use of parasitoids in biocontrol programs. Cotesia typhae nov. sp. (Fernández-Triana) (Hymenoptera: Braconidae) is a newly described parasitoid of the Mediterranean corn borer Sesamia nonagrioides (Lefebvre) (Lepidoptera: Noctuidae). Braconidae are known for their domesticated bracovirus, which is injected with eggs in the host larva to overcome its resistance. In this context, we compared reproductive success traits of four Kenyan strains of C. typhae on a French and a Kenyan populations of its host. Differences were found between the four strains and the two most contrasted ones were studied more thoroughly on the French host population. Parasitoid offspring size was correlated with parasitism success and the expression of bracovirus virulence genes (CrV1 and Cystatin) in the host larva after parasitism. Hybrids between these two parasitoid strains showed phenotype and gene expression profiles similar to the most successful parental strain, suggesting the involvement of dominant alleles in the reproductive traits. Ovary dissections revealed that the most successful strain injected more eggs in a single host larva than the less successful one, despite an equal initial ovocyte number in ovaries. It can be expected that the amount of viral particles increase with the number of eggs injected. The ability to bypass the resistance of the allopatric host may in consequence be related to the oviposition behaviour (eggs allocation). The influence of the number of injected eggs on parasitism success and on virulence gene expression was evaluated by oviposition interruption experiments.

Avoidance of host resistance in the oviposition-site preferences of rose bitterling

A contemporary outcome of dynamic host–parasite coevolution can be driven by the adaptation of a parasite to exploit its hosts at the population and species levels (parasite specialisation) or by local host adaptations leading to greater host resistance to sympatric parasite populations (host resistance). We tested the predominance of these two scenarios using cross-infection experiments with two geographically distant populations of the rose bitterling, Rhodeus ocellatus, a fish brood parasite of freshwater mussels, and four populations of their mussel hosts (two Anodonta woodiana and two Unio douglasiae populations) with varying degrees of geographic sympatry and local coexistence. Our data support predictions for host resistance at the species level but no effect of local coexistence between specific populations. Rhodeus ocellatus showed a preference for allopatric host populations, irrespective of host species. Host mussel response, in terms of ejection of R. ocellatus eggs, was stronger in the more widespread and abundant host species (A. woodiana) and this response tended to be higher in sympatric populations. These outcomes provide support for the importance of host resistance in bitterling oviposition-site decisions, demonstrating that host choice by R. ocellatus is adaptive by minimizing egg ejections. These findings imply that R. ocellatus, and potentially other bitterling species, may benefit from exploiting novel hosts, which may not possess appropriate adaptive responses to parasitism.

An experimental approach to the immuno-modulatory basis of host-parasite local adaptation in tapeworm-infected sticklebacks

The evolutionary arms race of hosts and parasites often results in adaptations, which may differ between populations. Investigation of such local adaptation becomes increasingly important to understand dynamics of host-parasite interactions and co-evolution. To this end we performed an infection experiment involving pairs of three-spined sticklebacks and their tapeworm parasite Schistocephalus solidus from three geographically separated origins (Germany, Spain and Iceland) in a fully-crossed design for sympatric and allopatric host/parasite combinations. We hypothesized that local adaptation of the hosts results in differences in parasite resistance with variation in parasite infection rates and leukocyte activation, whereas parasites from different origins might differ in virulence reflected in host exploitation rates (parasite indices) and S. solidus excretory-secretory products (SsESP) involved in immune manipulation.In our experimental infections, sticklebacks from Iceland were more resistant to S. solidus infection compared to Spanish and German sticklebacks. Higher resistance of Icelandic sticklebacks seemed to depend on adaptive immunity, whereas sticklebacks of German origin, which were more heavily afflicted by S. solidus, showed elevated activity of innate immune traits. German S. solidus were less successful in infecting and exploiting allopatric hosts compared to their Icelandic and Spanish conspecifics. Nevertheless, exclusively SsESP from German S. solidus triggered significant in vitro responses of leukocytes from naïve sticklebacks. Interestingly, parasite indices were almost identical across the sympatric combinations.Differences in host resistance and parasite virulence between the origins were most evident in allopatric combinations and were consistent within origin; i.e. Icelandic sticklebacks were more resistant and their S. solidus were more virulent in all allopatric combinations, whereas German sticklebacks were less resistant and their parasites less virulent. Despite such differences between origins, the degree of host exploitation was almost identical in the sympatric host-parasite combinations, suggesting that the local evolutionary arms race of hosts and parasites resulted in an optimal virulence, maximising parasite fitness while avoiding host overexploitation.