Local variation in the thistle species Cirsium grayanum affects its utilization by the herbivorous ladybird beetle Henosepilachna niponica

Host plants are the most critical environmental factor for phytophagous arthropods. Adaptation to a novel host will alter the distributional range of an herbivore to include the area the novel host covers, and might promote divergence between populations utilizing the old and new hosts. On the Oshima Peninsula, Hokkaido Island, northern Japan, the ladybird beetle Henosepilachna niponica (Lewis) (Coleoptera: Coccinellidae) usually occurs exclusively on the thistle Cirsium alpicola Nakai (Asteraceae), which is distributed in the southernmost part of the island, although some other Cirsium species often grow in sympatry. At least at one site (Assabu), however, H. niponica depends on another thistle, Cirsium grayanum (Maxim.) Nakai, the most abundant and widely distributed thistle species on the Oshima Peninsula. We examined adult feeding acceptance and preference, and larval performance, in two populations of H. niponica that utilize different species of thistles (the Assabu population occurs on C. grayanum; the Shiriuchi population, on C. alpicola) by testing three kinds of thistles (C. grayanum from Assabu, C. grayanum from Shiriuchi, and C. alpicola from Shiriuchi). Results strongly suggested that among the three plants tested, C. grayanum from Assabu was the most suitable host for H. niponica. We concluded that local differences in C. grayanum mainly explain why it is not used at Shiriuchi, but that there are also local differences in host use among the beetle populations, suggesting some degree of local adaptation. This suggests that different selective regimes associated with different host plants might promote divergence among beetle populations. We conclude that H. niponica is at present unlikely to expand its range to the whole range of C. grayanum.     

Does mother really know best? Oviposition preference reduces reproductive performance in the generalist parasitoid Aphidius ervi

The reproductive success of female parasitoids is dependent on their ability to accurately assess the suitability of a host for larval development. For generalist parasitoids, which utilize a broad range of species and instars as hosts, a set of assessment criteria determines whether a host is accepted or rejected. The suitability of a host, however, can only be imperfectly assessed by the female parasitoid, which can result in the selection of lesser quality hosts for oviposition. In this study we explored the disparity between host quality and host preference using the generalist koinobiotic parasitoid Aphidius ervi Haliday (Hymenoptera: Aphidiidae) and the host Aulacorthum solani (Harris) (Homoptera: Aphididae), the foxglove aphid. The second instar hosts produced the highest level of reproductive success, while third and fourth instars resulted in a substantially reduced reproductive performance. When given a choice of host instars, parasitoids preferred the older hosts for oviposition disregarding their reduced suitability for larval development. Results are discussed in context of mechanisms involved in A. ervi host selection and biases in the criteria used to assess hosts that may arise when parasitoids transfer host species between generations.     

Influence of rearing hosts on host size acceptance by Trichogramma australicum

We investigated the acceptance of different sized host models by Trichogramma australicum in the laboratory. We used isoline cultures of T. australicum reared in hosts of different sizes. Helicoverpa armigera represent relatively large hosts, and those of Sitotroga cerealella are small, termed the Ha and Sc biotypes, respectively. Five sizes of glass beads were tested for acceptance: diameter 0.5, 0.75, 1.0, 1.5 and 2.0 mm. The acceptance of a host model was determined by persistent attempted drilling of a glass bead by a female. The relationship between host egg size and number of eggs laid by a female was also investigated. We used three sizes of artificial egg (diameter 0.75, 1.00, and 1.50 mm of hemispherical cupules) each containing artificial diet. Ha biotype wasps accepted host models in the range 0.75--1.50 mm dia. (vol. 0.166--1.325 l), whereas those of the Sc biotype accepted host models in the range 0.50--1.00 mm dia. (vol. 0.05--0.393 l). This evidence suggests a lineal (possibly genetic) influence to host size acceptance for T. australicum, dependent on the size of the host in which the wasp has been reared. Also T. australicum lay fewer eggs in smaller artificial eggs than in larger ones. The role of host size in host acceptance and number of eggs delivered, and its implications, is discussed.     

Parasitism,host feeding and immature development of Encarsia formosa reared from Trialeurodes vaporariorum and Bemisia tabaci on Trialeurodes ricini

Encarsia formosa Gahan (Hymenoptera: Aphelinidae), a thelytokous parasitoid, is an important biological control agent of whiteflies because of its outstanding reproduction and host‐feeding ability. In this study, we evaluated the parasitism, host feeding and developmental time of E. formosa populations reared on Trialeurodes vaporariorum (Westwood) (EFT) or on Bemisia tabaci (Gennadius) (EFB) when different nymphal instars of the castor whitefly, Trialeurodes ricini (Misra), were offered as hosts, with an aim to understand the preference of the parasitoid on nymphal instars of T. ricini. Experiments were conducted on castor bean plants at 26 ± 2°C, 50–60% RH and 16 : 8 (L : D) photoperiod. The results showed that E. formosa successfully oviposited and fed on all nymphal instars of T. ricini. However, numbers of the first instars fed on by the E. formosa populations reared on T. vaporariorum (EFT) and B. tabaci (EFB) were significantly greater (45.9 and 31.3, respectively) than those of the second (EFT: 30.4 and EFB: 15.8), the third (EFT: 22.4 and EFB: 13.2) and the fourth nymphal instars (EFT: 6.0 and EFB: 3.8). The number of T. ricini nymphs parasitized by E. formosa varied significantly among different instars, and the parasitism rates on the first instar (EFT: 15.2; EFB: 7.7) and fourth instar (EFT: 19.3; EFB: 4.9) were greater than those on the second and third instars. Encarsia formosa reared on T. vaporariorum had a significantly higher host feeding and ovipositing potential on T. ricini than EFB. When parasitizing the fourth instar nymphs, E. formosa completed development in a significantly shorter time (12.9 day) than when ovipositing in other instars (17.8–19.1 day). These results showed that EFT had a better host adaption than EFB. The information from this study should be useful for us to better understand the performance and nymphal preference of E. formosa from T. vaporariorum and B. tabaci when they parasitized and fed on T. ricini, and the interactions of parasitoids with different host whitefly species.     

Factors determining the host plant range of the phytophagous mite, Tetranychus urticae (Acari: Tetranychidae): a method for quantifying host plant acceptance

The host plant acceptance of the phytophagous mite Tetranychus urticae was experimentally quantified. Host plant acceptance is described as the proportion of adult females settling on the test plant on which they have been placed. On the other hand, the host plant suitability of T. urticae on different plant species is expressed as the mean number of eggs produced by the females within 5 days (hereafter 'fecundity'). An inbred T. urticae line was tested with regard to host plant acceptance and fecundity on 11 potential host plants. These two variables were positively correlated across host plants; host plant species on which the fecundity was low were also those on which females settled less readily compared to host plants with high fecundity. The characteristics of host plant acceptance of the T. urticae are discussed in light of their potential food resource under natural conditions.     

Presence of two host races of Aphis gossypii Glover (Hemiptera: Aphididae) collected in Turkey

The aphid, Aphis gossypii, is a primary pest of citrus, cotton, cucurbits and greenhouse‐grown vegetables in Turkey and throughout Europe. There is some previous empirical data suggesting that host‐adapted genotypes of this aphid exist which may in fact be host‐races. To determine if host races of A. gossypii are indeed present in the eastern Mediterranean region of Turkey, reciprocal host transfer experiments and life table analyses were performed with multiple asexual lineages (= clones) of the aphid collected from different hosts. The collection hosts included citrus, cucumber, eggplant, okra, sweet pepper and cotton. Aphid developmental times on the host from which the aphid was originally collected (= collection or natal host) were shorter (5.2–6.0 days) and had a higher intrinsic rate of population growth (r_m = 0.25–0.44) than the 6.6–7.3 days required when the aphid was reared on a non‐original collection host (= non‐collection host or non‐natal host) and had r_m = 0.03–0.30. Total immature mortality of the cotton clone, especially in the first nymphal stage, was high (51–100%) with low r_m (0–0.03) on cucumber, citrus and sweet pepper. Aphid populations transferred from citrus, eggplant and okra to cotton (r_m = 0.29–0.30) did not differ significantly in their performance from that of the cotton population on cotton (r_m = 0.34), whereas that from sweet pepper and cucumber populations (r_m = 0.22–0.24) were significantly lower. These data have allowed us to separate A. gossypii into two distinct biological groups: (a) a ‘generalist’ population obtained from cucumber, sweet pepper, citrus, eggplant and okra which exhibited statistically better development on cotton; versus (b) a population from cotton which, by comparison on reciprocal hosts, developed poorly on non‐natal hosts except on eggplant. Development of the cotton clone on cucumber and okra was not improved after four successive generations on the non‐natal host. The good development of A. gossypii from eggplant and cotton on these reciprocal hosts suggests that these particular clones were similar, if not identical, host races.     

Host manipulation as a parasite transmission strategy when manipulation is exploited by non-host predators

Trophically transmitted parasites often alter their intermediate host's phenotype, thereby predisposing hosts to increased predation. This is generally considered to be a parasite strategy evolved to enhance transmission to the next host. However, the adaptive value of host manipulation is not clear, as it may be associated with costs, such as increased susceptibility to predator species that are unsuitable next hosts for the parasites. Thus, it has been proposed that, to be adaptive, manipulation should be specific by predisposing hosts more strongly to predation by target hosts (next host in the life cycle) than to non-hosts. Here we formally evaluate this prediction, and show that manipulation does not have to be specific to be adaptive. However, when manipulation is nonspecific, it needs to effectively increase the overall predation risk of infected hosts if it is to increase the parasite transmission probability. Thus, when initial predation risk is low, even highly nonspecific manipulation strategies can be adaptive. However, when initial predation risk is high, manipulation needs to be more specific to increase parasite transmission success. Therefore, nonspecific host manipulation may evolve in nature, but the adaptive value of a certain manipulation strategy can vary among different parasite populations depending on the variation in initial predation risk.     

The metazoan parasites of swordfish from Atlantic tropical–equatorial waters

The metazoan parasite fauna composition of the swordfish Xiphias gladius is reported from 145 (72 females and 73 males) specimens caught in Atlantic tropical–equatorial waters during June to July 2006 and August 2007. Seventeen metazoan taxa (two monogeneans, two digeneans, one acanthocephalan, four cestodes, seven nematodes and one copepod) were identified by morphological and genetic methodologies. Among those identified, Anisakis brevispiculata, Anisakis paggiae and the acanthocephalan Bolbosoma vasculosum are new host records. Dominant species were: the anisakid nematode larvae Anisakis physeteris, Anisakis brevispiculata, Hysterothylacium incurvum and Hysterothylacium corrugatum (s.l.), the adult cestode Fistulicola plicatus, the plerocercoid Hepatoxylon squali and the monogeneans Tristoma integrum and Tristoma coccineum, which showed high prevalences and intensities. The relationship between metazoan parasites and host size and sex was investigated for the first time. No statistically significant correlation was found between host sex and parasite burden. Infection levels with T. integrum and T. coccineum were observed to be significantly negatively correlated to host size. Infection levels by endoparasites observed in X.gladius from Atlantic tropical‐equatorial area may be related to host feeding behaviour and the availability of the parasites’ intermediate and final hosts in the area studied. Anisakids of the genus Anisakis, were identified genetically to species level, and were considered to be promising biological tags for the characterization of X.gladius stocks.     

Parasitic worms: strategies of host finding, recognition and invasion

Many parasitic worms enter their hosts by active invasion. Their transmission success is often based on a mass production of invasive stages. However, most stages show a highly specific host-finding behaviour. Information on host-finding mechanisms is available mainly for trematode miracidia and cercariae and for nematode hookworms. The larvae find and recognise their hosts, in some cases even with species specificity, via complex sequences of behavioural patterns with which they successively respond to various environmental and host cues. There is often a surprisingly high diversity of host-recognition strategies. Each parasite species finds and enters its host using a different series of cues. For example, different species of schistosomes enter the human skin using different recognition sequences. The various recognition strategies may reflect adaptations to distinct ecological conditions of transmission. Another question is how, after invasion, parasitic worms find their complex paths through their host's tissues to their often very specific microhabitats. Recent data show that the migrating parasite stages can follow local chemical gradients of skin and blood compounds, but their long-distance navigation within the host body still remains puzzling.

The high complexity, specificity and diversity of host-recognition strategies suggest that host finding and host recognition are important determinants in the evolution of parasite life cycles.     

Factors affecting the bionomics of the eastern African egg parasitoid Trichogramma bournieri Pintureau & Babault (Hymenoptera: Trichogrammatidae)

The trichogrammatid Trichogramma bournieri Pintureau & Babault is a polyphagous parasitoid of eggs of several cereal stemborer species in eastern Africa. The effects of host species, host age and duration of host deprivation on the performance of the parasitoid were studied in the laboratory. Host acceptance and suitability were tested using five stemborer species. The noctuids Sesamia calamistis Hampson, Sesamia nonagrioides (Lefebvre), Busseola fusca (Fuller) the crambid Chilo partellus (Swinhoe) and the pyralid Eldana saccharina Walker were successfully parasitized by T. bournieri. Parasitism, number of progeny and developmental time varied significantly with host species. The eggs of S. calamistis and B. fusca were the most suitable, whereas those of E. saccharina were the least suitable. While parasitism and number of progeny tended to decrease with age of hosts, there were no significant differences in sex ratio. Longevity of the parasitoid increased with increase in deprivation of hosts from 0 to 12 days. Average lifetime fecundity per female decreased, indicating resorption of eggs.     

Suitability of the maize weevil and angoumois grain moth as hosts for the parasitoidsAnisopteromalus calandrae andPteromalus cerealellae

Two parasitoids,Pteromalus cerealellae (Ashmead) andAnisopteromalus calandrae (Howard) (Hymenoptera: Pteromalidae), were compared for their ability to parasitize two important internally-developing insect pests of stored maize (Zea mays L.). Parasitism byP. cerealellae was greater on Angoumois grain moth,Sitotroga cerealella (Olivier), than on maize weevil,Sitophilus zeamais Motschulsky, in no-choice experiments.Anisopteromalus calandrae parasitized more maize weevils than didP. cerealellae. The former parasitoid parasitized only a few Angoumois grain moths successfully in maize, but parasitized many in wheat if the hosts were younger than 3 weeks old. Thus, both host age and type of grain affect suitability for parasitism. The effects of parental host (species on which the female developed) and experimental host (species exposed to parasitism) on parasitism rate ofP. cerealellae were tested in a host-switching experiment. Parasitism by parasitoids reared on maize weevils was 23% lower than that of parasitoids reared on Angoumois grain moth. This effect was independent of which host the filial generation of parasitoids was tested on. However, the experimental host species had a much greater effect on parasitoid fecundity than the parental host species. Female progeny had smaller body sizes when emerging from maize weevil than from Angoumois grain moth, which may explain the parental host effect on fecundity. There was also a slight intergenerational effect of host species on parasitoid body size.     

Ecological impacts of invading species: Do parasites of the cane toad imperil Australian frogs?

Parasite transfer to native fauna is a potentially catastrophic impact of invasive species. Introduced cane toads in Australia frequently host the nematode lungworm Rhabdias pseudosphaerocephala, which reduces viability of metamorph toads. If native frogs are vulnerable to this South American parasite, cane toad invasion may affect native species via this route; but if the native taxa are not vulnerable, we may be able to exploit the parasites for managing toads. Our laboratory experiments show that infective larvae can penetrate the body of all seven species of Australian frogs (five hylids: Cyclorana longipes, Litoria caerulea, Litoria dahlii, Litoria nasuta, Litoria rothii, one myobatrachid: Opisthodon ornatus, and one limnodynastid: Limnodynastes convexiusculus) we tested, but most did not host the adult worms at the end of the trials, and none showed major impairment of growth, survival or locomotor performance. One native tree‐frog (L. caerulea) retained high infection levels with few ill effects, suggesting that we might be able to use this taxon as a reservoir species to build up local parasite densities for toad management. However, the interspecific variation in lungworm retention suggests that generalizations about parasite effects on native frogs will be elusive.     

Divergent host plant specialization as the critical driving force in speciation between populations of a phytophagous ladybird beetle

Detecting the isolating barrier that arises earliest in speciation is critically important to understanding the mechanism of species formation. We tested isolating barriers between host races of a phytophagous ladybird beetle, Henosepilachna diekei (Coleoptera: Coccinellidae: Epilachnine), that occur sympatrically on distinct host plants. We conducted field surveys for the distribution of the beetles and host plants, rearing experiments to measure six potential isolating factors (adult host preference, adult and larval host performance, sexual isolation, egg hatchability, F(1) hybrid inviability, and sexual selection against F(1) hybrids), and molecular analyses of mitochondrial ND2 and the nuclear ITS2 sequences. We found significant genetic divergence between the host races, and extremely divergent host preference (i.e. habitat isolation) and host performance (i.e. immigrant inviability), but no other isolating barriers. The fidelity to particular host plants arises first and alone can prevent gene flow between differentiating populations of phytophagous specialists.     

Chemical signals mediating interactions betweenGaleruca tanaceti L. (Coleoptera, Chrysomelidae) and its egg parasitoidOomyzus galerucivorus (Hedqvits) (Hymenoptera, Eulophidae)

Chemical signals mediating interactions betweenGaleruca tanaceti and its egg parasitoidOomyzus galerucivorus (Hymenoptera, Eulophidae) were studied. Neither odor of gravid females ofG. tanaceti nor volatiles of their feces were attractive to the parasitoid. However, the presence of the beetles’ feces on a substrate arrested the parasitoid and elicited frequent antennal drumming. Thus, feces may contain a kairomone important for host finding. Odors of damaged and undamaged host plants had no effect on the parasitoids.O. galerucivorus did not detect its host eggs at close range but encountered them by chance. Neither the structure nor the dark color of the egg surface play a key role in host recognition, but chemicals of the extrachorion which could be isolated by dichloromethane. Fractionation of the dichloromethane extract by TLC revealed a single active fraction which induced host recognition behavior. Since the eggs ofG. tanaceti contain anthraquinones and anthrones which are active as feeding deterrents against predators, we hypothesized that reproductive success ofO. galerucivorus is due to sequestration of these protective compounds. However, GC-MS analyses revealed that there was no transfer of them from the host egg into the adult parasitoid.     

Sarcocystis rommeli,n. sp. (Apicomplexa: Sarcocystidae) from Cattle (Bos taurus) and its Differentiation from Sarcocystis hominis

Cattle (Bos taurus) are intermediate hosts for three named species of Sarcocystis, S. cruzi, S. hirsuta, and S. hominis. Recently, a fourth species was identified and named S. sinensis. However, S. sinensis originally named a species of Sarcocystis in water buffalo (Bubalus bubalis) in China. Based on unverifiable evidence, it was suggested that the same parasite infects cattle. In addition, S. sinensis was recently declared as nomen nudum because its naming violated the rules of International Code of Zoological Nomenclature. Thus, the fourth species using cattle as an intermediate host does not have a valid name. Here, we propose a new name, Sarcocystis rommeli for the S. sinensis‐like parasite from cattle in Argentina, and differentiate it ultrastructurally from S. hominis sarcocysts from experimentally infected cattle. Sarcocystis rommeli sarcocysts were microscopic with a 5‐μm‐thick wall with slender villar protrusions (Vp); the Vp were up to 5 μm long, up to 0.5 μm wide, and of uneven thickness, often bent at an angle. The ground substance layer (Gs) was up to 0.8 μm thick and smooth. Vesicular structures were seen at the base of the Vp. The bradyzoites were 10–12 μm long. Sarcocystis hominis sarcocysts had Vp that were often upright, up to 7.5 μm long, and up to 1.8 μm wide; the Gs was up to 2 μm thick and without vesicles. Its sarcocyst wall was up to 5.6 μm thick, the vp were bent at an angle, up to 5.8 μm long, the Gs was up to 2 μm thick, but without vesicles seen in S. rommeli. Beef containing sarcocysts of S. rommeli was not orally infectious for two human volunteers and a red fox (Vulpes vulpes). The Sarcocystis described here is molecularly different from S. cruzi, S. hirsuta, and S. hominis based on 18S rRNA and cox1 gene sequences.     

Effects of host stages and temperature on population parameters of Oomyzus sokolowskii, a larval-pupal parasitoid of Plutella xylostella

Oomyzus sokolowskii is alarval-pupal parasitoid of diamondback moth, Plutella xylostella. In a host stage preference test, the parasitoid parasitised all larval and pupal stages, but exhibited a strong preference for larvaeover prepupae or pupae, and did not show a preference among the larval instars. At 25°C, the developmental time, number and sex ratio of offspring per host pupa, and successful parasitism did not differ significantly among parasitoids reared from host larvae of different instars, indicating similar host suitability between larvae of different instars. Mean developmental times from egg to adult at 20, 22.5, 25, 30, 32.5, and 35°C were 26.5,21.0, 16.0, 12.7, 11.9 and 13.4 days, respectively. The favourable temperature range for development, survival, and reproduction of the parasitoid was 20--30°C. However, wasps that developed and emerged at a favourable temperature could parasitise effectively at 32--35°C for 24 hours. Life-fertility table studies at 20, 25, and 30°C showed that each female wasp on average parasitised 3.1, 13.2, 6.8 larvae of diamondback moth and produced 20.5, 92.1, 50.4 offspring, respectively, during her lifetime. The highest intrinsic rate of natural increase (r _m) of 0.263 female/day was reached at 30°C as a result of the short mean generation time at this temperature compared to that at 20 and 25°C, suggesting that the parasitoid had the highest potential for population growth at relatively high temperatures. This revised version was published online in July 2006 with corrections to the Cover Date.     

Effect of temperature on the pathogenicity of entomopathogenic nematodes (Steinernema and Heterorhabditis spp.) to Delia radicum

Susceptibility of last instarlarvae of Delia radicum to Steinernema feltiae, S. carpocapsae,S. arenarium, Heterorhabditismegidis and H. bacteriophora wasevaluated in the laboratory at 10 °C,15 °C and 20 °C. S. feltiaewas the only species that killed the larvae at10 °C; S. carpocapsae, S.arenarium and H. megidis were effectiveat 15–20 °C whereas H.bacteriophora killed the maggot only at20 °C. The temperature significantlyaffected the host searching ability of alltested species. Mobility was reduced at lowtemperatures. Significant effects of the hostpresence on nematode mobility were found forS. feltiae, S. arenarium and H. megidis but not for S. carpocapsaeand H. bacteriophora. The dynamics of theattachment to and penetration into the hostwere monitored for S. feltiae at10 °C, 15 °C or 20 °C and forS. carpocapsae at 20 °C. In theperiod of 6–30 hours after inoculation, S.carpocapsae attached in higher number at20 °C than did S. feltiae at alltemperatures. At 20 °C, S.carpocapsae penetrated the host only after 30hours while S. feltiae penetrated alreadyafter 15, 9, 6 hours at 10 °C,15 °C and 20 °C, respectively.     

Why resource history matters: age and oviposition history affect oviposition behaviour in exploiters of a mutualism

1. Acceptance of hosts for oviposition is often hardwired in short‐lived insects, but can be dynamic at the individual level due to variation in physiological state determinants such as ageing and prior oviposition. However, the effect of the oviposition history of resources together with time taken to accept less preferred hosts in ageing insects has rarely been investigated. 2. The time taken by parasitic fig wasps to accept resources with different oviposition histories was recorded in order to investigate the effect of wasp physiological state and resource oviposition history on oviposition behaviour. These wasps, which differ in life‐history traits, oviposit at specific developmental stages of enclosed fig inflorescences called syconia. 3. Behavioural assays were performed with naive wasps and wasps aged with and without prior oviposition experience. Syconia at the same developmental stage but differing in oviposition history were offered in no‐choice assays and the time taken to first oviposition attempt was recorded. 4. One short‐lived pro‐ovigenic galler species exhibited a decline with age in time taken to accept a syconium for oviposition. The exact timing of the transition from non‐acceptance to acceptance of less preferred syconia was determined in terms of the proportion of elapsed life span at the transition; this occurred at 25% of elapsed life span. 5. Longer‐lived parasitoids did not show any decline in specificity despite being aged for 50% of their life span. Therefore, host quality, trophic position, egg load and age may individually affect oviposition decisions or have interaction effects.