Infection patterns and new definitive host records for New Zealand gordiid hairworms (phylum Nematomorpha)

Some parasites modify the phenotype of their host in order to increase transmission to another host or to an environment suitable for reproduction. This phenomenon, known as host manipulation, is found across many parasite taxa. Freshwater hairworms are known for the behavioural changes they cause in their terrestrial arthropod hosts, increasing their likelihood of entering water to exit the host and reproduce. Understanding how infected arthropods move around in the natural environment could help uncover alterations in spatial distribution or movement induced by hairworms in their terrestrial definitive hosts. Moreover, few hairworm-host records exist for New Zealand, so any additional record could help elucidate their true host specificity. Here, we investigated whether infected terrestrial arthropods were more likely to approach streams in two subalpine communities of invertebrates, using a spatial grid of specialised pitfall traps. Although hairworm infection could not explain the movements of arthropod hosts near streams, we found several new host records for hairworms, including the first records for the recently described Gordionus maori. We also found some new host-parasite associations for mermithid nematodes. These records show that the host specificity of hairworms is quite low, suggesting that their diversity and distribution may be greater than what is currently known for New Zealand.     

Host preference and host suitability in an egg-pupal fruit fly parasitoid, Fopius arisanus (Sonan) (Hym., Braconidae)

The relative oviposition rate of the parasitoid Fopius arisanus (Sonan) was investigated across three frugivorous tephritid species, Bactrocera tryoni Froggart, Bactrocera jarvisi (Tryon) and Bactrocera cucumis French. Choice and no-choice tests were both used. The suitability of these three species for sustaining larval development and survival to the adult stage was also assessed. Fopius arisanus parasitized all three tephritid species, regardless of the method of exposure, but showed stronger preference for B. tryoni and B. jarvisi over B. cucumis. Superparasitism was extremely rare. Successful development of F. arisanus varied across host species. Bactrocera tryoni yielded significantly more parasitoids than B. jarvisi, but no wasps emerged from B. cucumis puparia. Tests were set up in replicated trials, but results were not homogeneous across trials. We discuss the host relationships of F. arisanus with reference to this variation and in relation to host suitability for larval development.     

Interkingdom host jumping underground: phylogenetic analysis of entomoparasitic fungi of the genus cordyceps

Most members of the ascomycetous genus Cordyceps are endoparasitic fungi of insects and other arthropods, but about 20 of the 300 described species are parasitic to hart's truffles, Elaphomyces spp. In order to understand the evolution of host specificity and the process of interkingdom host jumping in Cordyceps, we investigated the phylogenetic relationships of 22 representatives, including 4 truffle parasites and 18 insect parasites, based on nuclear and mitochondrial rDNA sequences. Five monophyletic groups were identified in both nuclear and mitochondrial phylogenies. In three of the five clades, the members utilized hosts from the same insect group, suggesting that the endoparasite-host connections have been conserved to some extent. On the other hand, it was also shown that major host shifts between distantly related insects must have occurred repeatedly. Notably, phylogenetic analyses strongly suggested that parasites of hart's truffles originated from parasites of cicada nymphs during the evolution of the CORDYCEPS: The common habitats of cicada nymphs and hart's truffles, deep underground and associated with tree roots, suggest that the interkingdom host jumping from Animalia to Fungi might have been promoted by the overlapping ecological niche of the unrelated hosts. This finding provides an impressive case of a drastic host shift in favor of the host habitat hypothesis.     

Divergent host acceptance behavior suggests host specialization in populations of the polyphagous mite Abacarus hystrix (Acari: Prostigmata: Eriophyidae)

For phytophagous arthropods, host acceptance behavior is a key character responsible for host plant specialization. The grain rust mite, Abacarus hystrix (Nalepa), is an obligately phytophagous, polyphagous eriophyid mite recorded from at least 70 grass species. In this study, the hypothesis that two host populations of this mite (one collected from quackgrass and the other from ryegrass) are highly host-specific was tested using behavioral data. For this purpose, female behavior when exposed to familiar and novel host plants was observed in no-choice cross experiments. Altogether, 13 variables were used to describe mite behavior. Data were subjected to principal component analysis, and host acceptance behavior was subsequently tested with generalized estimating equations (GEE). Distinct variation in female behavior between familiar and novel hosts was observed. Females from neither population accepted novel hosts. This was recorded as significant differences in the occupation of and overall activity on particular plant parts. On their familiar host, females were not active and showed little tendency to move. On novel hosts females were more active and mobile, spending more time walking, running, and climbing on the whole plant surface and showing a tendency to disperse. Other differences in behavior between studied populations were also observed. Thus, the results suggest that mites of these two studied populations (1) differ in their behaviors during plant exploitation and (2) can quickly distinguish between their familiar host and an unfamiliar host used by a conspecific. These findings support the hypothesis of narrow host specialization of ryegrass and quackgrass populations of this highly polyphagous species.     

Optimal foraging shapes host preference of a polyphagous leafminer

Abstract.  1. Most research on host selection by arthropods is based on optimal oviposition theory. This theory, also called the oviposition preference–offspring performance hypothesis, predicts that a female will choose those hosts for oviposition on which larvae perform best. Recent studies suggested, however, that optimal foraging by adults, or the quality of the host for adult performance, might also influence host choice.
2. This study investigated whether host preference of the polyphagous leafminer Liriomyza trifolii is determined by optimal foraging and/or optimal oviposition.
3. Female realised fecundity correlated nearly perfectly with feeding and oviposition preference of L. trifolii . The relationships between host preference and the offspring performance measures were always weaker. Hosts optimal for adult nutrition were also optimal for offspring performance but not vice versa . Hence, it was concluded that optimal foraging shapes feeding and oviposition preference of L. trifolii as this is the best strategy in order to maximise female fitness.     

Wolbachia host shifts: routes,mechanisms, constraints and evolutionary consequences

Wolbachia is one of the most abundant endosymbionts on earth, with a wide distribution especially in arthropods. Effective maternal transmission and the induction of various phenotypes in their hosts are two key features of this bacterium. Here, we review our current understanding of another central aspect of Wolbachia's success: their ability to switch from one host species to another. We build on the proposal that Wolbachia host shifts occur in four main steps: (i) physical transfer to a new species; (ii) proliferation within that host; (iii) successful maternal transmission; and (iv) spread within the host species. Host shift can fail at each of these steps, and the likelihood of ultimate success is influenced by many factors. Some stem from traits of Wolbachia (different strains have different abilities for host switching), others on host features such as genetic resemblance (e.g. host shifting is likely to be easier between closely related species), ecological connections (the donor and recipient host need to interact), or the resident microbiota. Host shifts have enabled Wolbachia to reach its enormous current incidence and global distribution among arthropods in an epidemiological process shaped by loss and acquisition events across host species. The ability of Wolbachia to transfer between species also forms the basis of ongoing endeavours to control pests and disease vectors, following artificial introduction into uninfected hosts such as mosquitoes. Throughout, we emphasise the many knowledge gaps in our understanding of Wolbachia host shifts, and question the effectiveness of current methodology to detect these events. We conclude by discussing an apparent paradox: how can Wolbachia maintain its ability to undergo host shifts given that its biology seems dominated by vertical transmission?     

Causal mechanisms underlying host specificity in bat ectoparasites

In parasites, host specificity may result either from restricted dispersal capacity or from fixed coevolutionary host-parasite adaptations. Knowledge of those proximal mechanisms leading to particular host specificity is fundamental to understand host-parasite interactions and potential coevolution of parasites and hosts. The relative importance of these two mechanisms was quantified through infection and cross-infection experiments using mites and bats as a model. Monospecific pools of parasitic mites (Spinturnix myoti and S. andegavinus) were subjected either to individual bats belonging to their traditional, native bat host species, or to another substitute host species within the same bat genus (Myotis). The two parasite species reacted differently to these treatments. S. myoti exhibited a clear preference for, and had a higher fitness on, its native host, Myotis myotis. In contrast, S. andegavinus showed no host choice, although its fitness was higher on its native host M. daubentoni. The causal mechanisms mediating host specificity can apparently differ within closely related host-parasite systems.     

How many species of arthropods? Erwin's estimate revised

Erwin's much debated estimate of 30 million species of arthropods is revised. The original estimate is based on the evaluation of host specificity of guilds in beetle samples, and subsequent hierarchical ratio extrapolations. The growing number of studies including mass sampling of arthropods have provided several data sets suitable for obtaining an empirical basis of this estimate. The structure in this modified version is somewhat changed compared to the original estimate in order to make each hierarchical step more easily testable. Plant species are separated into different growth forms, and host specificity measures are based only on phytophagous species. Effective specialization is applied as a measure of host specificity to correct for the fauna shared between plant species. A between community correction factor is applied to correct for differences in host specificity at different spatial scales. There are still great uncertainties attended with such estimates. The largest problems refer to the between community correction factor and the proportion of canopy species to total species. Further work on host specificity and the least known hyperdiverse groups are also needed. The revised version of the estimate does not support hyperestimates of 30 100 million species. Rather, it compares nicely with estimates derived from other estimation methods, indicating a global arthropod species richness of 5–10 million species.     

Influence of fruit traits on oviposition preference and offspring performance of Bactrocera tryoni (Froggatt) (Diptera: Tephritidae) on three tomato (Lycopersicon lycopersicum) cultivars

Abstract  In Queensland, three tomato ( Lycopersicon lycopersicum ) cultivars, Grosse Lisse, Roma and Cherry, are infested by Queensland fruit fly, Bactrocera tryoni (Froggatt). In this study, we examined if there was a correlation between oviposition preference and offspring performance of B. tryoni among the three tomato cultivars. We also investigated host plant traits that may explain any variation in preference and performance. Choice and no-choice experiments were carried out under laboratory conditions. A positive correlation between oviposition preference and offspring performance of B. tryoni was observed in the three tomato cultivars. Grosse Lisse and Roma cultivars were highly preferred by B. tryoni over Cherry cultivar. Performance (measured as proportion of eggs developing to the pupal stage) was significantly higher in Grosse Lisse and Roma cultivars than in Cherry cultivar. The pericarp toughness of Cherry cultivar appears responsible for its low rate of infestation, while the presence of 2-butanol and 1,4-butanediamine in Roma and Grosse Lisse, respectively, may partly be responsible for the high oviposition preference shown by B. tryoni towards these cultivars.     

Patterns of host specificity and transmission among parasites of wild primates

Multihost parasites have been implicated in the emergence of new diseases in humans and wildlife, yet little is known about factors that influence the host range of parasites in natural populations. We used a comprehensive data set of 415 micro- and macroparasites reported from 119 wild primate hosts to investigate broad patterns of host specificity. The majority (68%) of primate parasites were reported to infect multiple host species, including animals from multiple families or orders. This pattern corresponds to previous studies of parasites found in humans and domesticated animals. Within three parasite groups (viruses, protozoans and helminths), we examined parasite taxonomy and transmission strategy in relation to measures of host specificity. Relative to other parasite groups, helminths were associated with the greatest levels of host specificity, whereas most viruses were reported to infect hosts from multiple families or orders. Highly significant associations between the degree of host specificity and transmission strategy arose within each parasite group, but not always in the same direction, suggesting that unique constraints influence the host range of parasites within each taxonomic group. Finally characteristics of over 100 parasite species shared between wild primates and humans, including those recognised as emerging in humans, revealed that most of these shared parasites were reported from multiple host orders. Furthermore, nearly all viruses that were reported to infect both humans and non-human primates were classified as emerging in humans.     

Switch,disperse, repeat: host specificity is highly flexible in rodent-associated Eimeria

Interplay between conserved host specificity and occasional host switches is an important process determining the evolution of host-parasite systems. Here, we address the dynamics of host switches at the population level in rodent-associated Eimeria. Focusing mainly on two ecologically similar host groups, Murinae and Arvicolinae, we show that the Eimeria infecting those hosts form a complex system of many genetic lineages with different host specificities. The broad geographic distribution of lineages indicates that they are well-established genetic forms which retained their host specificities while spreading across large geographic areas. We also demonstrate that genetic structure is only partially reflected by morphological traits.     

Epiphyte host preferences and host traits: mechanisms for species-specific interactions

We investigated species-specific relationships among two species of vascular epiphytes and ten host tree species in a coastal plain forest in the southeastern United States. The epiphytes Tillandsia usneoides and Polypodium polypodioides were highly associated with particular host species in the field, but host traits that favored colonization were inadequate to fully explain the epiphyte-host associations for either epiphyte. Field transplant experiments that bypassed epiphyte colonization demonstrated that the growth of epiphytes was significantly higher on host tree species that naturally bore high epiphyte loads than on host species with few or no epiphytes. These species-specific relationships were highly correlated with the water-holding capacity of the host tree's bark. Positive and negative effects of throughfall, light attenuation by the canopy, and bark stability did not explain the overall patterns of host specificity, but did correlate with some epiphyte-host species relationships. The relative importance of particular host traits differed between the "atmospheric epiphyte" Tillandsia, and the fern Polypodium, which roots in the bark of its hosts. Species-specific interactions among plants, such as those described here, suggest that communities are more than individualistic assemblages of co-occurring species.     

Genetic delineation of sibling species of the pest fruit fly Bactocera (Diptera: Tephritidae) using microsatellites

Using a large set of microsatellites, the genetic relationships between three closely related Australian fruit fly species, Bactrocera tryoni (Froggatt), B. neohumeralis (Hardy) and B. aquilonis(May) were investigated. Bactrocera tryoni and B. neohumeralis are sympatric, while B. aquilonisis allopatric to both. The sympatric species, B. tryoni and B. neohumeralis, were found to be genetically distinct. It is likely that despite differences in mating time between these two species, some gene flow still occurs. In contrast, the sibling species B. tryoni and B. aquilonis were found to be closely related, despite allopatry. The level of genetic divergence was similar to that found within eastern Australian populations of B. tryoni. Consideration of all available genetic data suggests that this similarity is not due to recent (i.e. within the last 30 years) displacement of B. aquilonis by B. tryoni from the B. aquilonis region (north-western Australia). Instead the data suggests that, at least in the areas sampled, asymmetrical hybridization may have occurred over a longer timescale.     

Stand type is more important than red wood ant abundance for the structure of ground‐dwelling arthropod assemblages in managed boreal forests

• 1 The relationships between red wood ants (Formica rufa group) and other ground‐dwelling arthropods were studied in young managed forests stands in Eastern Finland. The main objectives were: (i) to test the influence of stand type (dominant tree species; age: sapling versus pole stage) and numbers of red wood ants on the occurrence of other ground‐dwelling arthropods and (ii) to study the occurrence of red wood ants versus other arthropods on a distance gradient from ant mounds. We used pitfall traps set in 5–14‐year‐old sapling stands and 30–45‐year‐old pole‐stage stands of Scots pine (Pinus sylvestris L.) and birch (Betula spp.) forests.
• 2 Pitfall trap catches of red wood ants did not vary significantly between the forest stand types, although some groups of other arthropods showed clear responses to stand type (e.g. catches of other Formicinae and Gnaphosidae were higher in sapling stands than in pole‐stage stands). The number of red wood ants clearly explained less of the variation in assemblages of other ground‐dwelling arthropods than the forest stand type.
• 3 Red wood ant numbers decreased significantly with distance from the mounds, but the other ground‐dwelling arthropods were insensitive to this gradient or even showed a preference for proximity to ant mounds and high ant activity.
• 4 The results obtained in the present study suggest that wood ants do not have strong effects on several other ground‐dwelling arthropod groups in young managed forests other than in the immediate vicinity of their mounds.

     

Coexistence of cytoplasmic incompatibility and male-killing-inducing endosymbionts, and their impact on host gene flow

Male-killing (MK) and cytoplasmic incompatibility (CI) inducing bacteria are among the most common endosymbionts of arthropods. Previous theoretical research has demonstrated that these two types of endosymbionts cannot stably coexist within a single unstructured host population if no doubly infected host individuals occur. Here, we analyse a model of two host subpopulations connected by migration. We demonstrate that coexistence of MK- and CI-inducing endosymbionts is possible if migration rates are sufficiently low. In particular, our results suggest that for coexistence to be possible, migration rates into the subpopulation infected predominantly with MK-inducing endosymbionts must be considerably low, while migration rates from the MK- to the CI-infected subpopulation can be very high. We also analyse how the presence of MK- and CI-inducing endosymbionts affects host gene flow between the two subpopulations. Employing the concept of the 'effective migration rate', we demonstrate that compared with an uninfected subdivided population, gene flow is increased towards the MK-infected island, but decreased towards the CI-infected island. We discuss our results with respect to the butterfly Hypolimnas bolina, in which infection polymorphism of CI- and MK-inducing Wolbachia has been reported across South-Pacific island populations.     

Adaptive learning of host preference in a herbivorous arthropod

Although many publications deal with the effects of experience on behaviour, adaptive learning (i.e. behavioural change with experience resulting in improved reproductive success) is poorly documented. We present direct evidence that learning of host preference improves fitness in the herbivorous mite, Tetranychus urticae . Individual mites from two strains were repeatedly given a choice between two host plants, tomato and cucumber, and then subjected to a performance test on each. For both strains, food experience affected the subsequent choice: individual mites learned to prefer cucumber over tomato. The performance test showed this effect to be adaptive, as the food plant the mites learned to prefer (cucumber) allowed for increased oviposition, survival and development. These findings have important implications for the interpretation of the preference–performance relationship among herbivorous arthropods. The frequently reported absence of such a relationship may be due to experience-dependent preference and/or performance.     

Geographical variation in host specificity of fleas (Siphonaptera) parasitic on small mammals: the influence of phylogeny and local environmental conditions

The evolution of host specificity remains a central issue in the study of host‐parasite relationships. Here we tackle three basic questions about host specificity using data on host use by fleas (Siphonaptera) from 21 geographical regions. First, are the host species exploited by a flea species no more than a random draw from the locally available host species, or do they form a taxonomically distinct subset? Using randomization tests, we showed that in the majority of cases, the taxonomic distinctness (measured as the average taxonomic distances among host species) of the hosts exploited by a flea is no different from that of random subsets of hosts taken from the regional pool. In the several cases where a difference was found, the taxonomic distinctness of the hosts used by a flea was almost always lower than that of the random subsets, suggesting that the parasites use hosts within a narrower taxonomic spectrum than what is available to them. Second, given the variation in host specificity among populations of the same flea species, is host specificity truly a species character? We found that host specificity measures are repeatable among different populations of the same flea species: host specificity varies significantly more among flea species than within flea species. This was true for both measures of host specificity used in the analyses: the number of host species exploited, and the index measuring the average taxonomic distinctness of the host species and its variance. Third, what causes geographical variation in host specificity among populations of the same flea species? In the vast majority of flea species, neither of our two measures of host specificity correlated with either the regional number of potential host species or their taxonomic distinctness, or the distance between the sampled region and the center of the flea's geographical range. However, in most flea species host specificity correlated with measures of the deviation in climatic conditions (precipitation and temperature) between the sampled region and the average conditions computed across the flea's entire range. Overall, these results suggest that host specificity in fleas is to a large extent phylogenetically constrained, while still strongly influenced by local environmental conditions.     

A comparative analysis of Wolbachia‐induced host reproductive phenotypes reveals transition rate heterogeneity

The endosymbiotic bacterium Wolbachia infects a wide range of arthropods and their relatives. It is an intracellular parasite transmitted through the egg from mother to offspring. Wolbachia can spread and persist through various means of host reproductive manipulation. How these different mechanisms of host manipulation evolved in Wolbachia is unclear. Which host reproductive phenotype is most likely to be ancestral and whether evolutionary transitions between some host phenotypes are more common than others remain unanswered questions. Recent studies have revealed multiple cases where the same Wolbachia strain can induce different reproductive phenotypes in different hosts, raising the question to what degree the induced host phenotype should be regarded as a trait of Wolbachia. In this study, we constructed a phylogenetic tree of Wolbachia and analyzed the patterns of host phenotypes along that tree. We were able to detect a phylogenetic signal of host phenotypes on the Wolbachia tree, indicating that the induced host phenotype can be regarded as a Wolbachia trait. However, we found no clear support for the previously stated hypothesis that cytoplasmic incompatibility is ancestral to Wolbachia in arthropods. Our analysis provides evidence for heterogeneous transition rates between host phenotypes.     

Are polyphagous geometrid moths with flightless females adapted to budburst phenology of local host species?

The majority of studies demonstrating local adaptation of insect herbivores involve sessile species, particularly those with a parthenogentic phase to their life history or endophagous "parasites" of plants. Current arguments suggest the strength of selection determines whether local adaptation can or cannot take place. Therefore local adaptation should not be limited to species with such traits. We studied the ability of three polyphagous geometrid moths with flightless adult females ( Erannis defoliaria , Operophtera brumata and O. fagata ) to synchronise their egg hatching with the budburst of a local host species in north east Scotland. A strong selection for hatching time is expected among generalist moths given the large variation in budburst phenology and an inability to hatch in synchrony with budburst decreases moth fitness substantially. In two successive seasons, we trapped emerging females from patches of five host species and recorded the temperature sum needed for 50% egg hatch of each brood laid by the trapped females. The hatching times of broods were compared against the average budburst time of the maternal host species in the study area. In addition, the trapping dates of each female were recorded. Only O. brumata showed synchrony with egg hatch and budburst which suggests local phenological adaptation to different host species . This could be maintained by selection and partial reproductive isolation between populations dwelling on different host species. No phenological adaptation was found in the other common geometrids of the study area.     

Biodiversity, host specificity, and dominance by eusocial species among sponge-dwelling alpheid shrimp on the Belize Barrier Reef

Alpheid shrimp represent an abundant and diverse, but poorly characterized, component of the cryptic biodiversity of coral reefs worldwide. Sponge‐inhabiting alpheids provide a promising model system for exploring patterns of cryptic reef biodiversity because their habitats (hosts) are discrete and qualitatively distinct units. We tabulated data from 14 years of collections at Carrie Bow Cay, Belize to quantify patterns of diversity, host specificity, and dominance among sponge‐dwelling shrimp (Synalpheus), with special attention to eusocial species. From > 600 sampled sponges of 17 species, we recognized at least 36 Synalpheus shrimp species. Of these, 15 (42%) were new to science. Species accumulation curves suggest that we have sampled most of the Synalpheus diversity at Carrie Bow Cay. Diversity of sponge‐dwelling Synalpheus was slightly higher in shallow water, probably because of greater habitat diversity, than in deep water. Host specificity was surprisingly high, with > 50% of all shrimp species found in only a single sponge species each, although some shrimp species used as many as six hosts. Cohabitation of individual sponges by multiple shrimp species was rarer than expected by chance, supporting previous distributional and behavioural evidence that competition for hosts is strong and moulds patterns of host association. The fauna of most well‐sampled sponge species was dominated, both in numbers of individuals and in frequency of occurrence, by eusocial species. Eusocial shrimp species also inhabited a significantly greater number of sponge species than did non‐social shrimp. Consequently, > 65% of shrimp in our quantitative samples belonged to the four eusocial species, and on a per‐species basis, eusocial species were 17 times as abundant as non‐social species. Our data suggest that the highly diverse sponge‐dwelling shrimp assemblage of the Belize Barrier Reef is structured by competition, and that eusociality has allowed a small number of species to dominate the sponge resource.