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In recent discussions, the widespread conviction that scientific individuation practices are governed by theories and concepts of biological individuality has been challenged, particularly by advocates of practice-based approaches. This discussion raises questions about the relationship between individuation practices and concepts of individuality. In this paper, I discuss four studies of host–parasite systems and analyze the respective individuation practices to see whether they correspond to established concepts of biological individuality. My analysis suggests that scientists individuate biological systems on different levels of organization and that the researchers' respective emphasis on one of the levels depends on the explanandum and research context as well as epistemic aims and purposes. It thus makes sense to use different concepts of individuality to account for different individuation practices. However, not all individuation practices are represented equally well by concepts of biological individuality. To account for this observation, I propose that concepts of individuality should be understood as abstracted, idealized, or simplified models that represent only certain aspects of scientific practice. A modeling account suggests a pluralistic view of concepts of biological individuality that not only allows the coexistence of different kinds of individuality (e.g., evolutionary individuality, immunological individuality, ecological individuality) but also of normative and descriptive concepts. [ABSTRACT FROM AUTHOR]

Climate change will alter interactions between parasites and their hosts. Warming may affect patterns of local adaptation, shifting the environment to favor the parasite or host and thus changing the prevalence of disease. We assessed local adaptation to hosts and temperature in the facultative ciliate parasite Lambornella clarki , which infects the western tree hole mosquito Aedes sierrensis. We conducted laboratory infection experiments with mosquito larvae and parasites collected from across a climate gradient, pairing sympatric or allopatric populations across three temperatures that were either matched or mismatched to the source environment. Lambornella clarki parasites were locally adapted to their hosts, with 2.6 times higher infection rates on sympatric populations compared with allopatric populations, but they were not locally adapted to temperature. Infection peaked at the intermediate temperature of 12.5°C, notably lower than the optimum temperature for free-living L. clarki growth, suggesting that the host's immune response can play a significant role in mediating the outcome of infection. Our results highlight the importance of host selective pressure on parasites, despite the impact of temperature on infection success. [ABSTRACT FROM AUTHOR]

Arms race dynamics are a common outcome of host–parasite coevolution. While they can theoretically be maintained indefinitely, realistic arms races are expected to be finite. Once an arms race has ended, for example due to the evolution of a generalist-resistant host, the system may transition into coevolutionary dynamics that favour long-term diversity. In microbial experiments, host–parasite arms races often transition into a stable coexistence of generalist-resistant hosts, (semi-)susceptible hosts, and parasites. While long-term host diversity is implicit in these cases, parasite diversity is usually overlooked. In this study, we examined parasite diversity after the end of an experimental arms race between a unicellular alga (Chlorella variabilis) and its lytic virus (PBCV-1). First, we isolated virus genotypes from multiple time points from two replicate microcosms. A time-shift experiment confirmed that the virus isolates had escalating host ranges, i.e., that arms races had occurred. We then examined the phenotypic and genetic diversity of virus isolates from the post-arms race phase. Post-arms race virus isolates had diverse host ranges, survival probabilities, and growth rates; they also clustered into distinct genetic groups. Importantly, host range diversity was maintained throughout the post-arms race phase, and the frequency of host range phenotypes fluctuated over time. We hypothesize that this dynamic polymorphism was maintained by a combination of fluctuating selection and demographic stochasticity. Together with previous work in prokaryotic systems, our results link experimental observations of arms races to natural observations of long-term host and parasite diversity. Graphical Abstract [ABSTRACT FROM AUTHOR]

Croziers paradox suggests that genetic kin recognition will not be evolutionarily stable. The problem is that more common tags (markers) are more likely to be recognized and helped. This causes common tags to increase in frequency, eliminating the genetic variability that is required for genetic kin recognition. Two potential solutions to this problem have been suggested: host-parasite coevolution and multiple social encounters. We show that the host-parasite coevolution hypothesis does not work as commonly assumed. Host-parasite coevolution only stabilizes kin recognition at a parasite resistance locus if parasites adapt rapidly to hosts and cause intermediate or high levels of damage (virulence). Additionally, when kin recognition is stabilized at a parasite resistance locus, this can have an additional cost of making hosts more susceptible to parasites. However, we show that if the genetic architecture is allowed to evolve, meaning natural selection can choose the recognition locus, genetic kin recognition is more likely to be stable. The reason for this is that host-parasite coevolution can maintain tag diversity at another (neutral) locus by genetic hitchhiking, allowing that other locus to be used for genetic kin recognition. These results suggest a way that host-parasite coevolution can resolve Croziers paradox, without making hosts more susceptible to parasites. However, the opportunity for multiple social encounters may provide a more robust resolution of Croziers paradox.

Introduction: Bottleneck events are crucial for the strength of genetic drift, selection and speed of evolution. They are believed to play a particularly prominent role for parasitic infrapopulations, inhabiting single host individuals, which are often established by very few parasite individuals during transmission. In vector-borne pathogens, the bottlenecking effects can even be serialized through repeated filtering of parasitic stages at different tissues and organs of the vector. Using qPCR we aimed to quantify the number of potentially transmittable sporozoites of the hemosporidian blood parasite Haemoproteus columbae in the specialized vector louse flies Pseudolynchia canariensis which transmit these parasites between house pigeon hosts Columba livia. Results: Based on qPCR measurements of organ-derived DNA of individual louse flies, we estimate that the midgut of these vectors contains on average 20 parasites, the hindgut and other intestines ca. 50 parasites and the salivary glands ca. 5 parasite cells. Nearly one third of all vector individuals appeared to lack parasite DNA, despite having only infected hosts as blood meal sources. The magnitude of parasite numbers in midgut and salivary glands tended to correlate positively. Discussion: Our results indicate, potential severe bottlenecking of parasite populations during individual transmission events and a probable effect of individual vector immunity on this variable. However, this may be partly alleviated by the coloniality of house pigeons, the frequency of louse flies and their daily feeding events in most populations, leading to repeated transmission opportunities, decreased quasi-vertical transmission between parents and offspring and probable panmixia of Haemoproteus columbae lineages. Many of these mechanisms might not apply in other host-vector systems. We propose several additional molecular and microscopical tools to improve the accuracy of estimating parasite population sizes in vectors and call for more estimations in different vector species to better understand the co-evolution between malarialike blood parasites and their avian and insect hosts. [ABSTRACT FROM AUTHOR]

Host–parasite systems, including social parasites that exploit resources of the host colonies, are fascinating objects for evolutionary biologists mainly due to the dynamic and often rapid host–parasite coevolution. Host‐switching events are believed to induce rapid speciation of parasitic species. The socially parasitic ant lineage Myrmoxenus, which corresponds to the monophyletic Temnothorax corsicus group, counts in total a dozen species. Most Myrmoxenus species utilize a single host species, but a few others, like Myrmoxenus ravouxi (André, 1896) and M. gordiagini Ruzsky, 1902, are known to use multiple host taxa. Myrmoxenus zaleskyi (Sadil, 1953) was described as a putative congener of M. ravouxi based on its distinct host selection. In this paper, we investigate the diversity of the widely distributed European lineages M. ravouxi and M. zaleskyi from multiple and complementary perspectives to understand whether the host preference exhibited by these two forms implies speciation. We integrated evidence from molecular genetics using mitochondrial CO I/CO II genes, including the tLeu‐region, and multivariate analyses of morphometric data collected from workers and female sexuals (gynes). Although there is substantial regional host species specificity, results suggest that host switching did not result in phylogenetic or morphological divergence and that the central European M. zaleskyi can be considered the junior synonym of M. ravouxi. As the lineage Myrmoxenus has been the subject of considerable evolutionary research, these results are essential to achieve a more accurate picture of host–parasite systems in the future and further strengthen the justification of an integrative approach in studying similarly complex systems. We advise against describing new parasitic species based on host preference unless coupled with marked heritable phenotypic adaptations. [ABSTRACT FROM AUTHOR]

Previous host-parasite coevolutionary theory has focused on understanding the determinants of local adaptation using spatially discrete models. However, these studies fall short of describing patterns of host-parasite local adaptation across spatial scales. In contrast, empirical work demonstrates that patterns of adaptation depend on the scale at which they are measured. Here, we propose a mathematical model of host-parasite coevolution in continuous space that naturally leads to a scale-dependent definition of local adaptation. In agreement with empirical research, we find that patterns of adaptation vary across spatial scales. In some cases, not only the magnitude of local adaptation but also the identity of the locally adapted species will depend on the spatial scale at which measurements are taken. Building on our results, we suggest a way to consistently measure parasite local adaptation when continuous space is the driver of cross-scale variation. We also describe a way to test whether continuous space is driving cross-scale variation. Taken together, our results provide a new perspective that can be used to understand empirical observations previously unexplained by theoretical expectations and deepens our understanding of the mechanics of host-parasite local adaptation. [ABSTRACT FROM AUTHOR]

The high tree diversity in tropical forests has long been a puzzle to ecologists. In the 1970s, Janzen and Connell proposed that tree species (hosts) coexist due to the stabilizing actions of specialized enemies. This Janzen–Connell hypothesis was subsequently supported by theoretical studies. Yet, such studies have taken the presence of specialized pathogens for granted, overlooking that pathogen coexistence also requires an explanation. Moreover, stable ecological coexistence does not necessarily imply evolutionary stability. What are the conditions that allow Janzen–Connell effects to evolve? We link theory from community ecology, evolutionary biology and epidemiology to tackle this question, structuring our approach around five theoretical frameworks. Phenomenological Lotka–Volterra competition models provide the most basic framework, which can be restructured to include (single‐ or multi‐)pathogen dynamics. This ecological foundation can be extended to include pathogen evolution. Hosts, of course, may also evolve, and we introduce a coevolutionary model, showing that host–pathogen coevolution can lead to highly diverse systems. Our work unpacks the assumptions underpinning Janzen–Connell and places theoretical bounds on pathogen and host ecology and evolution. The five theoretical frameworks taken together provide a stronger theoretical basis for Janzen–Connell, delivering a wider lens that can yield important insights into the maintenance of diversity in these increasingly threatened systems. [ABSTRACT FROM AUTHOR]

IntroductionBottleneck events are crucial for the strength of genetic drift, selection and speed of evolution. They are believed to play a particularly prominent role for parasitic infrapopulations, inhabiting single host individuals, which are often established by very few parasite individuals during transmission. In vector-borne pathogens, the bottlenecking effects can even be serialized through repeated filtering of parasitic stages at different tissues and organs of the vector. Using qPCR we aimed to quantify the number of potentially transmittable sporozoites of the hemosporidian blood parasite Haemoproteus columbae in the specialized vector louse flies Pseudolynchia canariensis which transmit these parasites between house pigeon hosts Columba livia.ResultsBased on qPCR measurements of organ-derived DNA of individual louse flies, we estimate that the midgut of these vectors contains on average 20 parasites, the hindgut and other intestines ca. 50 parasites and the salivary glands ca. 5 parasite cells. Nearly one third of all vector individuals appeared to lack parasite DNA, despite having only infected hosts as blood meal sources. The magnitude of parasite numbers in midgut and salivary glands tended to correlate positively.DiscussionOur results indicate, potential severe bottlenecking of parasite populations during individual transmission events and a probable effect of individual vector immunity on this variable. However, this may be partly alleviated by the coloniality of house pigeons, the frequency of louse flies and their daily feeding events in most populations, leading to repeated transmission opportunities, decreased quasi-vertical transmission between parents and offspring and probable panmixia of Haemoproteus columbae lineages. Many of these mechanisms might not apply in other host-vector systems. We propose several additional molecular and microscopical tools to improve the accuracy of estimating parasite population sizes in vectors and call for more estimations in different vector species to better understand the co-evolution between malaria-like blood parasites and their avian and insect hosts.

Crozier’s paradox suggests that genetic kin recognition will not be evolutionarily stable. The problem is that more common tags (markers) are more likely to be recognized and helped. This causes common tags to increase in frequency, eliminating the genetic variability that is required for genetic kin recognition. Two potential solutions to this problem have been suggested: host–parasite coevolution and multiple social encounters. We show that the host–parasite coevolution hypothesis does not work as commonly assumed. Host–parasite coevolution only stabilizes kin recognition at a parasite resistance locus if parasites adapt rapidly to hosts and cause intermediate or high levels of damage (virulence). Additionally, when kin recognition is stabilized at a parasite resistance locus, this can have an additional cost of making hosts more susceptible to parasites. However, we show that if the genetic architecture is allowed to evolve, meaning natural selection can choose the recognition locus, genetic kin recognition is more likely to be stable. The reason for this is that host–parasite coevolution can maintain tag diversity at another (neutral) locus by genetic hitchhiking, allowing that other locus to be used for genetic kin recognition. These results suggest a way that host–parasite coevolution can resolve Crozier’s paradox, without making hosts more susceptible to parasites. However, the opportunity for multiple social encounters may provide a more robust resolution of Crozier’s paradox. [ABSTRACT FROM AUTHOR]

Using a dynamic optimisation model for juvenile fish in stochastic food environments, we investigate optimal hormonal regulation, energy allocation and foraging behaviour of a growing host infected by a parasite that only incurs an energetic cost. We find it optimal for the infected host to have higher levels of orexin, growth and thyroid hormones, resulting in higher activity levels, increased foraging and faster growth. This growth strategy thus displays several of the fingerprints often associated with parasite manipulation: higher levels of metabolic hormones, faster growth, higher allocation to reserves (i.e. parasite‐induced gigantism), higher risk‐taking and eventually higher predation rate. However, there is no route for manipulation in our model, so these changes reflect adaptive host compensatory responses. Interestingly, several of these changes also increase the fitness of the parasite. Our results call for caution when interpreting observations of gigantism or risky host behaviours as parasite manipulation without further testing. [ABSTRACT FROM AUTHOR]

Abstract Background Understanding the interactions between bat flies and host bats offer us fundamental insights into the coevolutionary and ecological processes in host-parasite relationships. Here, we investigated the identities, host specificity, and patterns of host association of bat flies in a subtropical region in East Asia, which is an understudied region for bat fly research. Methods We used both morphological characteristics and DNA barcoding to identify the bat fly species found on 11 cavernicolous bat species from five bat families inhabiting Hong Kong. We first determined the phylogenetic relationships among bat fly species. Then, we elucidated the patterns of bat-bat fly associations and calculated the host specificity of each bat fly species. Furthermore, we assembled the mitogenomes of three bat fly species from two families (Nycteribiidae and Streblidae) to contribute to the limited bat fly genetic resources available. Results We examined 641 individuals of bat flies and found 20 species, of which many appeared to be new to science. Species of Nycteribiidae included five Nycteribia spp., three Penicillidia spp., two Phthiridium spp., one Basilia sp., and one species from a hitherto unknown genus, whereas Streblidae included Brachytarsina amboinensis, three Raymondia spp., and four additional Brachytarsina spp. Our bat-bat fly association network shows that certain closely related bat flies within Nycteribiidae and Streblidae only parasitized host bat species that are phylogenetically more closely related. For example, congenerics of Raymondia only parasitized hosts in Rhinolophus and Hipposideros, which are in two closely related families in Rhinolophoidea, but not other distantly related co-roosting species. A wide spectrum of host specificity of these bat fly species was also revealed, with some bat fly species being strictly monoxenous, e.g. nycteribiid Nycteribia sp. A, Phthiridium sp. A, and streblid Raymondia sp. A, while streblid B. amboinensis is polyxenous. Conclusions The bat fly diversity and specificity uncovered in this study have shed light on the complex bat-bat fly ecology in the region, but more bat-parasite association studies are still needed in East Asian regions like China as a huge number of unknown species likely exists. We highly recommend the use of DNA barcoding to support morphological identification to reveal accurate host-ectoparasite relationships for future studies. Graphical Abstract

Social parasites usually rely on chemical cues (cuticular hydrocarbons) to successfully invade and coexist with their hosts. Most ants that are obligate social parasites (slave-makers) can parasitize several related host species with different levels of chemical similarity although there are few exceptions where there is only a single host species. An example of the latter is Rossomyrmex minuchae, which was known to be only associated with Proformica longiseta. However, a recent discovery of a new R. minuchae population revealed that it can parasitize P. nasuta, a species with a separate distribution to P. longiseta. Chemical analyses of cuticular hydrocarbons show local adaptation of the parasite to its host in this new population, being more similar to P. nasuta than to other R. minuchae populations. In addition, genetic analyses evidence differences from the other known populations, from which it split 1.21 Mya during glacial and interglacial periods of the Pleistocene and remained separated to the present day. This historical genetic isolation and the chemical differences found between parasite populations may evidence a speciation process and support the local host–parasite coevolution. [ABSTRACT FROM AUTHOR]

Abstract Using a dynamic optimisation model for juvenile fish in stochastic food environments, we investigate optimal hormonal regulation, energy allocation and foraging behaviour of a growing host infected by a parasite that only incurs an energetic cost. We find it optimal for the infected host to have higher levels of orexin, growth and thyroid hormones, resulting in higher activity levels, increased foraging and faster growth. This growth strategy thus displays several of the fingerprints often associated with parasite manipulation: higher levels of metabolic hormones, faster growth, higher allocation to reserves (i.e. parasite‐induced gigantism), higher risk‐taking and eventually higher predation rate. However, there is no route for manipulation in our model, so these changes reflect adaptive host compensatory responses. Interestingly, several of these changes also increase the fitness of the parasite. Our results call for caution when interpreting observations of gigantism or risky host behaviours as parasite manipulation without further testing.

Monogenea (Platyhelminthes), mainly gill and fin ectoparasites of fish, are often recognized as host specific and morphologically and ecologically diverse. These parasites exhibit high species diversity at the level of host species or individual fish specimens. Using case studies, especially those widely performed in Dactylogyrus parasitizing cyprinoid fish, this article presents current knowledge on the ecology and evolution of congeneric gill monogeneans. The important aspects of the ecology of congeneric monogeneans are highlighted, in particular: host specificity expressed at several host levels (from strict specificity to phylogenetic specificity), microhabitat specificity expressed by restricted positions on fish gills to facilitate intraspecific mating, and the link between microhabitat preference and morphological adaptation (i.e., sclerotized structures of the haptor) or reproductive isolation. From the evolutionary perspective, this study focused on the processes of the speciation and diversification of congeneric monogeneans, highlighting the role of host switch as the most prominent coevolutionary event, accompanied in some cases mostly by intrahost speciation or cospeciation, as revealed by cophylogenetic studies. Here, important knowledge on evolutionary patterns of host specificity, microhabitat specificity, and morphological adaptation is presented. Host-specific monogeneans may represent an important tool for studying the historical biogeography of their hosts. Specifically, in the case of freshwater fish hosts exhibiting disjunctive distribution, they reflect both historical and contemporary contacts. The role of host-specific congeneric monogeneans in revealing historical intercontinental and intracontinental contacts between freshwater fish is highlighted. Finally, the importance of the role of genetic coadaptation, limiting the presence of host-specific monogeneans in hybrid fish, is emphasized.

Background: Understanding the interactions between bat flies and host bats offer us fundamental insights into the coevolutionary and ecological processes in host-parasite relationships. Here, we investigated the identities, host specificity, and patterns of host association of bat flies in a subtropical region in East Asia, which is an understudied region for bat fly research. Methods: We used both morphological characteristics and DNA barcoding to identify the bat fly species found on 11 cavernicolous bat species from five bat families inhabiting Hong Kong. We first determined the phylogenetic relationships among bat fly species. Then, we elucidated the patterns of bat-bat fly associations and calculated the host specificity of each bat fly species. Furthermore, we assembled the mitogenomes of three bat fly species from two families (Nycteribiidae and Streblidae) to contribute to the limited bat fly genetic resources available. Results: We examined 641 individuals of bat flies and found 20 species, of which many appeared to be new to science. Species of Nycteribiidae included five Nycteribia spp., three Penicillidia spp., two Phthiridium spp., one Basilia sp., and one species from a hitherto unknown genus, whereas Streblidae included Brachytarsina amboinensis, three Raymondia spp., and four additional Brachytarsina spp. Our bat-bat fly association network shows that certain closely related bat flies within Nycteribiidae and Streblidae only parasitized host bat species that are phylogenetically more closely related. For example, congenerics of Raymondia only parasitized hosts in Rhinolophus and Hipposideros, which are in two closely related families in Rhinolophoidea, but not other distantly related co-roosting species. A wide spectrum of host specificity of these bat fly species was also revealed, with some bat fly species being strictly monoxenous, e.g. nycteribiid Nycteribia sp. A, Phthiridium sp. A, and streblid Raymondia sp. A, while streblid B. amboinensis is polyxenous. Conclusions: The bat fly diversity and specificity uncovered in this study have shed light on the complex bat-bat fly ecology in the region, but more bat-parasite association studies are still needed in East Asian regions like China as a huge number of unknown species likely exists. We highly recommend the use of DNA barcoding to support morphological identification to reveal accurate host-ectoparasite relationships for future studies. [ABSTRACT FROM AUTHOR]

Background: Several hypotheses have been proposed to explain parasite infection in parental species and their hybrids. Hybrid heterosis is generally applied to explain the advantage for F1 generations of hybrids exhibiting a lower level of parasite infection when compared to parental species. Post-F1 generations often suffer from genetic incompatibilities potentially reflected in the higher level of parasite infection when compared to parental species. However, the presence of specific parasites in an associated host is also limited by close coevolutionary genetic host-parasite associations. This study focused on monogenean parasites closely associated with two leuciscid fish species—common bream and roach—with the aim of comparing the level of monogenean infection between parental species and hybrids representing two F1 generations with different mtDNA and two backcross generations with different cyto-nuclear compositions. Results: Monogenean infection in F1 generations of hybrids was lower when compared to parental species, in line with the hybrid heterosis hypothesis. Monogenean infection in backcross generations exhibited similarities with the parental species whose genes contributed more to the backcross genotype. The distribution of monogeneans associated with one or the other parental species showed the same asymmetry with a higher proportion of roach-associated monogeneans in both F1 generations and backcross generation with roach in the paternal position. A higher proportion of common bream-associated monogeneans was found in backcross generation with common bream in the paternal position. Conclusions: Our study indicated that cyto-nuclear incompatibilities in hybrids do not induce higher monogenean infection in backcross generations when compared to parental species. However, as backcross hybrids with a higher proportion of the genes of one parental taxon also exhibited high level of this parental taxon-associated parasites, host-parasite coevolutionary interactions seem to play an obvious role in determining the level of infection of host-specific monogeneans in hybrids. [ABSTRACT FROM AUTHOR]

George Price showed how the effects of natural selection and environmental change could be mathematically partitioned. This partitioning may be especially useful for understanding host–parasite coevolution, where each species represents the environment for the other species. Here, we use coupled Price equations to study this kind of antagonistic coevolution. We made the common assumption that parasites must genetically match their host's genotype to avoid detection by the host's self/nonself recognition system, but we allowed for the possibility that non‐matching parasites have some fitness. Our results show how natural selection on one species results in environmental change for the other species. Numerical iterations of the model show that these environmental changes can periodically exceed the changes in mean fitness due to natural selection, as suggested by R.A. Fisher. Taken together, the results give an algebraic dissection of the eco‐evolutionary feedbacks created during host–parasite coevolution. [ABSTRACT FROM AUTHOR]

Hosts can avoid parasites (and pathogens) by reducing social contact, but such isolation may carry costs, e.g. increased vulnerability to predators. Thus, many predator–host–parasite systems confront hosts with a trade-off between predation and parasitism. Parasites, meanwhile, evolve higher virulence in response to increased host sociality and consequently, increased multiple infections. How does predation shift coevolution of host behaviour and parasite virulence? What if predators are selective, i.e. predators disproportionately capture the sickest hosts? We answer these questions with an eco-coevolutionary model parametrized for a Trinidadian guppy–Gyrodactylus spp. system. Here, increased predation drives host coevolution of higher grouping, which selects for higher virulence. Additionally, higher predator selectivity drives the contact rate higher and virulence lower. Finally, we show how predation and selectivity can have very different impacts on host density and prevalence depending on whether hosts or parasites evolve, or both. For example, higher predator selectivity led to lower prevalence with no evolution or only parasite evolution but higher prevalence with host evolution or coevolution. These findings inform our understanding of diverse systems in which host behavioural responses to predation may lead to increased prevalence and virulence of parasites. [ABSTRACT FROM AUTHOR]

Abstract George Price showed how the effects of natural selection and environmental change could be mathematically partitioned. This partitioning may be especially useful for understanding host–parasite coevolution, where each species represents the environment for the other species. Here, we use coupled Price equations to study this kind of antagonistic coevolution. We made the common assumption that parasites must genetically match their host's genotype to avoid detection by the host's self/nonself recognition system, but we allowed for the possibility that non‐matching parasites have some fitness. Our results show how natural selection on one species results in environmental change for the other species. Numerical iterations of the model show that these environmental changes can periodically exceed the changes in mean fitness due to natural selection, as suggested by R.A. Fisher. Taken together, the results give an algebraic dissection of the eco‐evolutionary feedbacks created during host–parasite coevolution.

Obligate brood parasitic birds lay their eggs in the nests of other species, reducing the host's own reproductive output. To circumvent these fitness costs, many—but not all—host species have evolved the ability to recognize and reject brood parasitic eggs. What factors constrain egg rejection, and why do host species vary in their likelihood of rejection? Previous comparative studies have found that egg rejection rates covary with several biotic factors (including larger body size, smaller relative brain size, and more northerly breeding latitudes), but much behavioral variation in the occurrence of egg rejection remains unexplained. In this study, we test a corollary of the maternal investment hypothesis, by assessing whether species with higher clutch sizes are more likely to eliminate parasitic eggs. We examined two published data sets comprising over 200 unique bird species, controlling for phylogeny and other known interspecific correlates of egg rejection rates. Contrary to the prediction, we found no evidence for a positive relationship between clutch size and egg rejection rate. Rather, our analyses suggest a weak but consistent negative relationship between absolute and relative metrics of clutch size versus egg rejection rate across species. These results are instead consistent with two previously proposed alternative hypotheses: that egg rejection is constrained by a trade-off between maternal investment and anti-parasitic defenses, possibly mediated by endocrine mechanisms linked to parental care, and/or that cognitive decision rules facilitate the detection of dissimilar eggs in smaller clutches. [ABSTRACT FROM AUTHOR]

Obligate parasites need one or more hosts to complete their life cycle. However, hosts might show intraspecific variation in quality with respect to the parasites themselves, thus affecting on‐host and off‐host parasite performance. High heritability in host quality for the parasite may therefore exert long‐lasting selective pressures on the parasite and influence host–parasite coevolution. However, the amount of variation and heritability in host quality are unknown for most parasite species, especially in wild populations of hosts. Both measures were estimated in a wild‐caught bird Parus major that was experimentally infested by two developmental stages (larva and nymph) of a ectoparasite (the tick Ixodes arboricola). We examined variation in host quality through variation in tick performance, namely the on‐host performance (attachment success, feeding time, engorgement weight and feeding success) and the off‐host performance (moulting time, moulting success and overall survival). Herein we also investigated the influence on tick performance of host traits linked with the bird's life history and physiology such as body condition, sex, age and haematocrit. By correlating tick performance variables between larvae and nymphs feeding on the same bird at different times, we found a significant correlation in attachment success, suggesting consistent among‐host variation for this performance measure, but no significant larva‐nymph correlations for the other tick variables. Animal models relating tick performance variables to the host pedigree showed a strong heritable signal for host quality as measured through tick feeding time, and lower but substantial estimates in other performance variables. With regard to the host traits, feeding success and survival of tick larvae were lower on female birds, and nymphal survival was higher on older birds. Larval feeding time was negatively correlated with host haematocrit. This is one of the first studies showing consistent intraspecific variation and heritability of host quality for a multi‐stage ectoparasite. [ABSTRACT FROM AUTHOR]

The dynamics of coevolution between hosts and parasites are influenced by their genetic interactions. Highly specific interactions, where the outcome of an infection depends on the precise combination of host and parasite genotypes (G × G interactions), have the potential to maintain genetic variation by inducing negative frequency‐dependent selection. The importance of this effect also rests on whether such interactions are consistent across different environments or modified by environmental variation (G × G × E interaction). In the black bean aphid, Aphis fabae, resistance to its parasitoid Lysiphlebus fabarum is largely determined by the possession of a heritable bacterial endosymbiont, Hamiltonella defensa, with strong G × G interactions between H. defensa and L. fabarum. A key environmental factor in this system is the host plant on which the aphid feeds. Here, we exposed genetically identical aphids harbouring three different strains of H. defensa to three asexual genotypes of L. fabarum and measured parasitism success on three common host plants of A. fabae, namely Vicia faba, Chenopodium album and Beta vulgaris. As expected, we observed the pervasive G × G interaction between H. defensa and L. fabarum, but despite strong main effects of the host plants on average rates of parasitism, this interaction was not altered significantly by the host plant environment (no G × G × E interaction). The symbiont‐conferred specificity of resistance is thus likely to mediate the coevolution of A. fabae and L. fabarum, even when played out across diverse host plants of the aphid. [ABSTRACT FROM AUTHOR]

Natural host populations differ in their susceptibility to infection by parasites, and these intrapopulation differences are still an incompletely understood component of host‐parasite dynamics. In this study, we used controlled infection experiments with wild‐caught guppies (Poecilia reticulata) and their ectoparasite Gyrodactylus turnbulli to investigate the roles of local adaptation and host genetic composition (immunogenetic and neutral) in explaining differences in susceptibility to infection. We found differences between our four study host populations that were consistent between two parasite source populations, with no indication of local adaptation by either host or parasite at two tested spatial scales. Greater values of host population genetic variability metrics broadly aligned with lower population mean infection intensity, with the best alignments associated with major histocompatibility complex (MHC) "supertypes". Controlling for intrapopulation differences and potential inbreeding variance, we found a significant negative relationship between individual‐level functional MHC variability and infection: fish carrying more MHC supertypes experienced infections of lower severity, with limited evidence for supertype‐specific effects. We conclude that population‐level differences in host infection susceptibility probably reflect variation in parasite selective pressure and/or host evolutionary potential, underpinned by functional immunogenetic variation. [ABSTRACT FROM AUTHOR]

Social interactions with conspecifics are key to the fitness of most animals and, through the transmission opportunities they provide, are also key to the fitness of their parasites. As a result, research to date has largely focused on the role of host social behavior in imposing selection on parasites, particularly their virulence and transmission phenotypes. However, host social behavior also influences the distribution of parasites among hosts, with implications for their evolution through non-random mating, gene flow, and genetic drift, and thus ability to respond to that selection. Here, we review the paucity of empirical studies on parasites, and draw from empirical studies of free-living organisms and population genetic theory to propose several mechanisms by which host social behavior potentially drives parasite evolution through these less-well studied mechanisms. We focus on the guppy host and Gyrodactylus (Monogenea) ectoparasitic flatworm system and follow a spatially hierarchical outline to highlight that social behavior varies between individuals, and between host populations across the landscape, generating a mosaic of ecological and evolutionary outcomes for their infecting parasites. We argue that the guppy-Gyrodactylus system presents a unique opportunity to address this fundamental knowledge gap in our understanding of the connection between host social behavior and parasite evolution. Individual differences in host social behavior generates fine-scale changes in the spatial distribution of parasite genotypes, shape the size, and diversity of their infecting parasite populations and may generate non-random mating on, and non-random transmission between hosts. While at population and metapopulation level, variation in host social behavior interacts with landscape structure to affect parasite gene flow, effective population size, and genetic drift to alter the coevolutionary potential of local adaptation. Significance statement: Social interactions between animals shape the evolution of the pathogens that infect them. Most research exploring this phenomenon has focused on the selection such interactions impose, but social hosts also shape parasite evolution by determining the ability of their parasites to respond to that selection. Here, we explore how host social behavior drives parasite evolution by shaping non-random mating, gene flow, and genetic drift, from the scale of the individual to the landscape. The relative strength of these evolutionary mechanisms can have striking implications for the evolution of parasite traits such as virulence and alter the evolutionary trajectories of populations across the landscape. We emphasize the importance of studies combining parasite population genetics, host social behavior, and landscape processes to illuminate complex host-parasite coevolutionary dynamics. [ABSTRACT FROM AUTHOR]

Three subspecies of Northern Bahamian Rock Iguanas, Cyclura cychlura , are currently recognized: C. c. cychlura, restricted to Andros Island, and C. c. figginsi and C. c. inornata, native to the Exuma Island chain. Populations on Andros are genetically distinct from Exuma Island populations, yet genetic divergence among populations in the Exumas is inconsistent with the 2 currently recognized subspecies from those islands. The potential consequences of this discrepancy might include the recognition of a single subspecies throughout the Exumas rather than 2. That inference also ignores evidence that populations of C. cychlura are potentially adaptively divergent. We compared patterns of population relatedness in a three-tiered host-parasite system: C. cychlura iguanas, their ticks (genus Amblyomma , preferentially parasitizing these reptiles), and Rickettsia spp. endosymbionts (within tick ectoparasites). Our results indicate that while C. c. cychlura on Andros is consistently supported as a separate clade, patterns of relatedness among populations of C. c. figginsi and C. c. inornata within the Exuma Island chain are more complex. The distribution of the hosts, different tick species, and Rickettsia spp. supports the evolutionary independence of C. c. inornata. Further, these patterns are also consistent with two independent evolutionarily significant units within C. c. figginsi. Our findings suggest coevolutionary relationships between the reptile hosts, their ectoparasites, and rickettsial organisms, suggesting local adaptation. This work also speaks to the limitations of using neutral molecular markers from a single focal taxon as the sole currency for recognizing evolutionary novelty in populations of endangered species. [ABSTRACT FROM AUTHOR]

Trichomonosis caused by the flagellate Trichomonas gallinae is one of the most important avian diseases worldwide. The parasite is localised in the oesophageal area of its host and mainly infects pigeon and dove species. During the last decade, a host expansion to passerine birds occurred, making the disease a potential threat for passerine predators as naïve host species. Here, we investigated the effect of the parasite on two Accipiter species in Germany which show a comparable lifestyle but differ in prey choice, the Northern goshawk (Accipiter gentilis) mainly hunting pigeons and the Eurasian sparrowhawk (Accipiter nisus) mainly feeding on passerines. We genetically identified the parasite strains using the Fe-Hydrogenase gene as marker locus and compared the incidence of parasite presence and clinical signs of trichomonosis between nestlings of the two Accipiter species. In total, we identified 14 strains, with nine strains unknown so far. There was a higher strain diversity and prevalence of Trichomonas spp. in goshawks than sparrowhawks (42.4% vs. 21.2%) whereas sparrowhawks when being infected more often displayed clinical signs of trichomonosis than goshawks (37.1% vs. 6.1%). Even though sparrowhawks were mainly infected with the finch epidemic strain and genetic data indicated some variation between isolates, no correlation with virulence could be detected. All in all, goshawks seem to be better adapted to Trichomonas infections, whereas to sparrowhawks, this is a novel disease with more severe manifestations, from individual morbidity to a higher risk of population decline caused by trichomonosis. [ABSTRACT FROM AUTHOR]

Understanding the transition of bacterial species from commensal to pathogen, or vice versa, is a key application of evolutionary theory to preventative medicine. This requires working knowledge of the molecular interaction between hosts and bacteria, ecological interactions among microbes, spatial variation in bacterial prevalence or host life history, and evolution in response to these factors. However, there are very few systems for which such broad datasets are available. One exception is the gram-negative bacterium, Helicobacter pylori, which infects upwards of 50% of the global human population. This bacterium is associated with a wide breadth of human gastrointestinal disease, including numerous cancers, inflammatory disorders, and pathogenic infections, but is also known to confer fitness benefits to its host both indirectly, through interactions with other pathogens, and directly. Outstanding questions are therefore why, when, and how this bacterium transitions along the parasitism-mutualism continuum. We examine known virulence factors, genetic predispositions of the host, and environmental contributors that impact progression of clinical disease and help define geographical trends in disease incidence. We also highlight the complexity of the interaction and discuss future therapeutic strategies for disease management and public health in light of the longstanding evolutionary history between the bacterium and its human host.

Host‐parasite coevolution is ubiquitous, shaping genetic and phenotypic diversity and the evolutionary trajectory of interacting species. With the advances of high throughput sequencing technologies applicable to model and non‐model organisms alike, it is now feasible to study in greater detail (a) the genetic underpinnings of coevolution, (b) the speed and type of dynamics at coevolving loci, and (c) the genomic consequences of coevolution. This review focuses on three recently developed approaches that leverage information from host and parasite full genome data simultaneously to pinpoint coevolving loci and draw inference on the coevolutionary history. First, co‐genome‐wide association study (co‐GWAS) methods allow pinpointing the loci underlying host‐parasite interactions. These methods focus on detecting associations between genetic variants and the outcome of experimental infection tests or on correlations between genomes of naturally infected hosts and their infecting parasites. Second, extensions to population genomics methods can detect genes under coevolution and infer the coevolutionary history, such as fitness costs. Third, correlations between host and parasite population size in time are indicative of coevolution, and polymorphism levels across independent spatially distributed populations of hosts and parasites can reveal coevolutionary loci and infer coevolutionary history. We describe the principles of these three approaches and discuss their advantages and limitations based on coevolutionary theory. We present recommendations for their application to various host (prokaryotes, fungi, plants, and animals) and parasite (viruses, bacteria, fungi, and macroparasites) species. We conclude by pointing out methodological and theoretical gaps to be filled to extract maximum information from full genome data and thereby to shed light on the molecular underpinnings of coevolution. [ABSTRACT FROM AUTHOR]

Pathogens are generally expected to evolve faster than their hosts and are therefore likely to be locally adapted. However, some pathogens might lag behind in the co-evolutionary arms race because they do not have some of the advantages shared by most other pathogens (e.g., high mutation or recombination rates, short generation time, high dispersal ability). This is the case of Microbotryum fungi that cause the anther smut disease in plants of the family Caryophyllaceae. We investigated the patterns of local adaptation and maladaptation in Microbotryum carthusianorum and its host plant Dianthus carthusianorum. We performed a full cross-inoculation experiment using half-sib plant families and fungal samples originating from three naturally infected populations in the Czech Republic. We specifically asked, which components of pathogen fitness (i.e., infectivity and host manipulation) are affected by local (mal)adaptation. The pathogen was on average 1.6 times more successful in infecting plants from foreign populations compared to plants from its home population. Once the infection was successful, the pathogen accelerated the plant's flowering and thus increased the opportunity for transmission to new hosts. However, the level of manipulation of host flowering did not differ between home and foreign populations. This study showed that the pathogen's infectivity followed a clear pattern of local maladaptation, whereas the host manipulation did not. Our study taken together with previous studies of a related anther smut species reveals the pervasiveness of local maladaptation in this group of pathogens that arises as the result of their restricted gene flow and reduced recombination rates. [ABSTRACT FROM AUTHOR]

Host–parasite dynamics often vary over time, brought about by changes in the parasite's virulence or the host's ability to resist or tolerate the parasite. Although virulence evolution in microparasites is well studied, we know little about temporal change in the pathogenicity of macroparasites such as blood‐feeding insects. Using data collected over 35 yr, we report a reduction in pathogenicity of the hematophagous swallow bug (Cimex vicarius) on its Cliff Swallow (Petrochelidon pyrrhonota) host. Relative to experimentally fumigated, parasite‐free nests, the negative effects of bugs on nestling swallow body mass and survival were less in the later years of the study than in the earlier years, and the negative relationship between nestling body mass and bug abundance became weaker over time. The survival of adult birds exposed to swallow bugs increased throughout the study, while survival of birds from parasite‐free nests decreased over time. Swallow bug abundance per nest, bug body size, and bug age ratios did not change during the study. Between‐colony transmission of bugs showed less immigration into larger colonies than in earlier studies, but there was no net change in transmission. Cliff Swallows did not reduce their exposure to bugs over time by being more likely to avoid infested nest or colony sites. Parents increased the number of food deliveries to their offspring over time in the presence of parasites, but the total amount of food delivered was unchanged. The reduced cost of swallow bug ectoparasitism does not seem related to changes in parasite narrow‐sense virulence, the host's avoidance of parasites, the presence of alternative hosts for bugs, or climate‐driven phenological mismatches. The results probably reflect the Cliff Swallow's evolving of greater tolerance to swallow bugs, brought about by the bird's shift from natural cliff nesting sites to artificial structures that may harbor more bugs than natural cliffs. This study shows that hosts can respond relatively rapidly to high levels of parasitism, and provides support for models that suggest the evolution of tolerance should be expected in some host–parasite systems. [ABSTRACT FROM AUTHOR]

Abstract The ectoparasitic mite, Varroa destructor, shifted host from the eastern honeybee, Apis cerana, to the western honeybee, Apis mellifera. Whereas the original host survives infestations by this parasite, they are lethal to colonies of its new host. Here, we investigated a population of A. cerana naturally infested by the V. destructor Korea haplotype that gave rise to the globally invasive mite lineage. Our aim was to better characterize traits that allow for the survival of the original host to infestations by this particular mite haplotype. A known major trait of resistance is the lack of mite reproduction on worker brood in A. cerana. We show that this trait is neither due to a lack of host attractiveness nor of reproduction initiation by the parasite. However, successful mite reproduction was prevented by abnormal host development. Adult A. cerana workers recognized this state and removed hosts and parasites, which greatly affected the fitness of the parasite. These results confirm and complete previous observations of brood susceptibility to infestation in other honeybee host populations, provide new insights into the coevolution between hosts and parasites in this system, and may contribute to mitigating the large‐scale colony losses of A. mellifera due to V. destructor.

Two types of Apis cerana are known from India: black or hill bees and yellow or plains bees, which differ in mitochondrial haplotype. We investigated whether black and yellow A. cerana host different species or strains of Varroa. Apis. cerana and any associated Varroa mites were collected from 21 colonies (5 with Varroa), in Bengaluru, India, and surrounding areas. Bees were scored for color and mitochondrial haplotype using sequences of the cytochrome oxidase I and II genes and the intervening leucine tRNA gene and non-coding region. Mites were scored for species and strain using partial sequence of cytochrome oxidase I. We found both V. destructor with a haplotype similar to the previously reported Sri Lanka1 and a novel haplotype of V. jacobsoni. Contrary to our expectations, we found no association of mite species and bee mitochondrial haplotype. [ABSTRACT FROM AUTHOR]

Major histocompatibility complex (MHC)-based mating rules can evolve as a way to avoid inbreeding or to increase offspring immune competence. While the role of mating preference in shaping the MHC diversity in vertebrates has been acknowledged, its impact on individual MHC diversity has not been considered. Here, we use computer simulations to investigate how simple mating rules favouring MHC-dissimilar partners affect the evolution of the number of MHC variants in individual genomes, accompanying selection for resistance to parasites. We showed that the effect of such preferences could sometimes be dramatic. If preferences are aimed at avoiding identical alleles, the equilibrium number of MHC alleles is much smaller than under random mating. However, if the mating rule minimizes the ratio of shared to different alleles in partners, MHC number is higher than under random mating. Additionally, our simulations revealed that a negative correlation between the numbers of MHC variants in mated individuals can arise from simple rules of MHC-disassortative mating. Our results reveal unexpected potential of MHC-based mating preferences to drive MHC gene family expansions or contractions and highlight the need to study the mechanistic basis of such preferences, which is currently poorly understood. [ABSTRACT FROM AUTHOR]

Granulins are conserved in nearly all metazoans, with an intriguing loss in insects. These pleiotropic peptides are involved in numerous physiological and pathological processes yet have been overwhelmingly examined in mammalian systems. While work in other animal models has been informative, a richer understanding of the proteins should be obtained by integrating knowledge from all available contexts. The main bodies of work described here include 1) the structure-function relationships of progranulin and its cleavage products, 2) the role of expanded granulin gene families and different isoforms in fish immunology, 3) the release of granulin peptides to promote host angiogenesis by parasitic worms, 4) a diversity of molluscan uses for granulins, including immune activation in intermediate hosts to trematodes, 5) knowledge gained on lysosomal functions from C. elegans and the stress-related activities of granulins. We provide an overview of functional reports across the Metazoa to inform much-needed future research. • Granulins are conserved and pleiotropic, yet the evolution of functions is unclear. • Functional patterns and novelties across animal granulins are detailed. • The loss of granulins in some insects is addressed. • Granulin can be released from parasites and hosts simultaneously. • Future directions for each type of organism are suggested throughout. [ABSTRACT FROM AUTHOR]

The initial process by which novel sexual signals evolve remains unclear, because rare new variants are susceptible to loss by drift or counterselection imposed by prevailing female preferences. 1,2,3,4 We describe the diversification of an acoustic male courtship signal in Hawaiian populations of the field cricket Teleogryllus oceanicus , which was brought about by the evolution of a brachypterous wing morph ("small-wing") only 6 years ago. 5 Small-wing has a genetic basis and causes silence or reduced-amplitude signaling by miniaturizing male forewings, conferring protection against an eavesdropping parasitoid, Ormia ochracea. 5 We found that wing reduction notably increases the fundamental frequency of courtship song from an average of 5.1 kHz to 6.4 kHz. It also de-canalizes male song, broadening the range of peak signal frequencies well outside normal song character space. As courtship song prompts female mounting and is sexually selected, 6,7,8,9 we evaluated two scenarios to test the fate of these new signal values. Females might show reduced acceptance of small-wing males, imposing counterselection via prevailing preferences. Alternatively, females might accept small-wing males as readily as long-wing males if their window of preference is sufficiently wide. Our results support the latter. Females preferred males who produced some signal over none, but they mounted sound-producing small-wing males as often as sound-producing long-wing males. Indiscriminate mating can facilitate the persistence of rare, novel signal values. If female permissiveness is a general characteristic of the earliest stages of sexual signal evolution, then taxa with low female mate acceptance thresholds should be more prone to diversification via sexual selection. [Display omitted] • Sexual signal evolution requires that novel traits escape loss due to selection • In Hawaiian crickets, small wings evolved several years ago, diversifying male song • Females mate with singing small-wing males as readily as with long-wing males • Permissive female mating can facilitate rapid sexual trait diversification Zhang et al. test how new animal signal variants evolve. In Hawaiian crickets, males evolved higher-pitch courtship songs under selection from eavesdropping enemies. However, males singing songs at these abnormal frequencies suffer no mating disadvantage. Permissive females may buffer against the loss of new signal variants. [ABSTRACT FROM AUTHOR]

Abstract Background Animal and plant species can harbour microbes that provide them with protection against enemies. These beneficial microbes can be a significant component of host defence that complement or replaces a repertoire of immunity, but they can also be costly. Given their impact on host and parasite fitness, defensive microbes have the potential to influence host-parasite interactions on an evolutionary timescale. Results Using a phenotypic framework, we explore the evolutionary and coevolutionary dynamics of a host-parasite interaction in the presence of defensive microbes. We show that costs of host-defensive microbe systems are critical in determining whether a defensive microbe based system or an immune system provides better host protection investment. Partitioning the coevolutionary dynamics yields testable predictions. The density of defensive microbes influences the strength of selection resulting from host - defensive microbe - parasite coevolutionary interactions. We find that they lessen the negative effects of infection on hosts and reduce infectivity by directly competing with parasites. Conclusions Defensive microbes might thus play a central role in host-parasite interactions, by outright replacing host-based defences, engaging in within-host competition with parasites, and ultimately driving tripartite coevolutionary dynamics.

The cost of males should give asexual females an advantage when in competition with sexual females. In addition, high‐fecundity asexual genotypes should have an advantage over low‐fecundity clones, leading to reduction in clonal diversity over time. To evaluate fitness components in a natural population, we measured the annual reproductive rate of individual sexual and asexual female Potamopyrgus antipodarum, a New Zealand freshwater snail, in field enclosures that excluded competitors and predators. We used allozyme genotyping to assign the asexual females to particular clonal genotypes. We found that the most fecund asexual clones had similar or higher fecundity as the top 10% of sexual families, suggesting that fecundity selection, even without the cost of males, would lead to replacement of the sexual population by clones. Consequently, we expected that the clones with the highest fecundity would dominate the natural population. Counter to this prediction, we found that high annual reproductive rates did not correlate with the frequency of clones in the natural population. When we exposed the same clones to parasites in the laboratory, we found that resistance to infection was positively correlated with the frequency of clones in the population. The correlation between fecundity and parasite resistance was negative, suggesting a trade‐off between these two traits. Our results thus suggest that parasite resistance is an important short‐term predictor of the success of asexual P. antipodarum in this population. [ABSTRACT FROM AUTHOR]

Parasites often jump to and become established in a new host species. There is much evidence that the probability of such host shifts decreases with increasing phylogenetic distance between donor and recipient hosts, but the consequences of such preferential host switching remain little explored. We develop a computational model to investigate the dynamics of parasite host shifts in the presence of this phylogenetic distance effect. In this model, a clade of parasites evolves on an evolving clade of host species where parasites can cospeciate with their hosts, switch to new hosts, speciate within hosts or become extinct. Our model predicts that host phylogenies are major determinants of parasite distributions across trees. In particular, we predict that trees consisting of few large clades of host species and those with fast species turnover should harbor more parasites than trees with many small clades and those that diversify more slowly. Within trees, large clades are predicted to exhibit a higher fraction of infected species than small clades. We discuss our results in the light of recent cophylogenetic studies in a wide range of host–parasite systems. [ABSTRACT FROM AUTHOR]

CRISPR-Cas is an adaptive prokaryotic immune system that prevents phage infection. By incorporating phage-derived 'spacer' sequences into CRISPR loci on the host genome, future infections from the same phage genotype can be recognized and the phage genome cleaved. However, the phage can escape CRISPR degradation by mutating the sequence targeted by the spacer, allowing them to re-infect previously CRISPR-immune hosts, and theoretically leading to coevolution. Previous studies have shown that phage can persist over long periods in populations of Streptococcus thermophilus that can acquire CRISPR-Cas immunity, but it has remained less clear whether this coexistence was owing to coevolution, and if so, what type of coevolutionary dynamics were involved. In this study, we performed highly replicated serial transfer experiments over 30 days with S. thermophilus and a lytic phage. Using a combination of phenotypic and genotypic data, we show that CRISPR-mediated resistance and phage infectivity coevolved over time following an arms race dynamic, and that asymmetry between phage infectivity and host resistance within this system eventually causes phage extinction. This work provides further insight into the way CRISPR-Cas systems shape the population and coevolutionary dynamics of bacteria-phage interactions. [ABSTRACT FROM AUTHOR]

Ant nests are relatively stable and long-lasting microhabitats that attract a diverse group of arthropods. Particular stressors, however, can trigger ants to relocate their nest to a new site. It is unclear how associated arthropods respond to occasional nest moving of their host. Here, I report field observations which showed that the potentially parasitic larvae of the beetle Clytra quadripunctata follow their red wood ant host during nest relocation, either by crawling on their own or by being carried by the host workers. These observations shed new light on the spatial dynamics between ants and their associates. [ABSTRACT FROM AUTHOR]

Abstract The coevolution of parasites and their hosts is accompanied by rapid reciprocal adaptations. Recent studies focusing on the evolutionary ecology of parasitic fungi and their host insects have revealed an unexpectedly complex molecular arms race. Insects recognize the presence of invading fungi when molecular patterns associated with the fungal cell wall bind to host receptors and respond by inducing the synthesis of proteinase inhibitors and antifungal peptides such as metchnikowin. Parasitic fungi can in turn sense the presence of these host-derived defense molecules and respond by synthesizing chymotrypsin and metalloproteases to digest them. As a counter-response, insects have evolved the ability to perceive the presence of microbial metalloproteinases, inducing the synthesis of further antimicrobial peptides and the insect metalloproteinase inhibitor to regulate these virulence-associated fungal proteinases. These specific responses and counter-responses indicate that coevolution has resulted in communication between fungal parasites and their insect hosts, which was recently shown to involve transcriptional reprogramming controlled by histone acetylation/deacetylation and DNA methylation. Consequently, the reciprocal adaptations associated with host–parasite coevolution can be expanded beyond the genetic level to encompass epigenetic mechanisms. [ABSTRACT FROM AUTHOR]

The genetic diversity of Varroa destructor (Anderson & Trueman) is limited outside its natural range due to population bottlenecks and its propensity to inbreed. In light of the arms race between V. destructor and its honeybee (Apis mellifera L.) host, any mechanism enhancing population admixture of the mite may be favored. One way that admixture can occur is when two genetically dissimilar mites coinvade a brood cell, with the progeny of the foundresses admixing. We determined the relatedness of 393 pairs of V. destructor foundresses, each pair collected from a single bee brood cell (n = five colonies). We used six microsatellites to identify the genotypes of mites coinvading a cell and calculated the frequency of pairs with different or the same genotypes. We found no deviation from random coinvasion, but the frequency of cells infested by mites with different genotypes was high. This rate of recombination, coupled with a high transmission rate of mites, homogenized the allelic pool of mites within the apiary. [ABSTRACT FROM AUTHOR]

Bats are important zoonotic reservoirs for many pathogens worldwide. Although their highly specialized ectoparasites, bat flies (Diptera: Hippoboscoidea), can transmit Bartonella bacteria including human pathogens, their eco-epidemiology is unexplored. Here, we analyzed the prevalence and diversity of Bartonella strains sampled from 10 bat fly species from 14 European bat species. We found high prevalence of Bartonella spp. in most bat fly species with wide geographical distribution. Bat species explained most of the variance in Bartonella distribution with the highest prevalence of infected flies recorded in species living in dense groups exclusively in caves. Bat gender but not bat fly gender was also an important factor with the more mobile male bats giving more opportunity for the ectoparasites to access several host individuals. We detected high diversity of Bartonella strains (18 sequences, 7 genotypes, in 9 bat fly species) comparable with tropical assemblages of bat-bat fly association. Most genotypes are novel (15 out of 18 recorded strains have a similarity of 92-99%, with three sequences having 100% similarity to Bartonella spp. sequences deposited in GenBank) with currently unknown pathogenicity; however, 4 of these sequences are similar (up to 92% sequence similarity) to Bartonella spp. with known zoonotic potential. The high prevalence and diversity of Bartonella spp. suggests a long shared evolution of these bacteria with bat flies and bats providing excellent study targets for the eco-epidemiology of host-vector-pathogen cycles. [ABSTRACT FROM AUTHOR]

Studies on interactions between Orobanche species and their hosts are mainly focused on resistance, radical interactions, and haustorium development. The influence of the host plant on morphology of the parasite under environmental controlled conditions is not reported in the literature. With the aim of filling this gap, the results of cultivation experiments of Orobanche crenata on Vicia faba, V. sativa, Pisum sativum, Cicer arietinum, and Lathyrus clymenum are presented. The tested population of O. crenata infected all the studied hosts but showed preferences toward Vicia and Pisum and, in particular, toward a foreign V. sativa cv. The higher resistance of the evaluated local cv. could be due to coevolution between host and parasite. There was little variation in times of emergence: about 110 days on V. faba, while around 125 on P. sativum, C. arietinum, and V. sativa. The shape and size of flowers of O. crenata were influenced by its hosts. The more flourishing the host, the more vigorous the parasite with respect to overall size, number of flowers, and density of the inflorescence. However, significant variations were also observed in the shape of the calyx and the colour of the corolla and stigma. These observations suggest that other taxa of Orobanche that grow in the wild, which are considered to be different species, could be morphotypes of the same taxon, although a process of speciation may be in progress. [ABSTRACT FROM AUTHOR]