Using the literature to evaluate parasitoid host ranges: a case study of Macrocentrus grandii (Hymenoptera: Braconidae) introduced into North America to control Ostrinia nubilalis (Lepidoptera: Crambidae)

We propose a method for using the literature to evaluate host ranges of parasitoids that are candidates for biological control introductions. Data on the parasitoids that attack a given host species can be used as negative evidence concerning the candidate whose host range is being evaluated. By compiling studies for a variety of host species, one can delineate those taxa unlikely to be attacked by the candidate. Using a retrospective case study of a parasitoid introduced into North America, we describe (1) this approach to using the literature to evaluate host range and (2) how well predictions based on such an evaluation match actual host range. Based on the host range of Macrocentrus grandii in Eurasia as reported in the literature, we predicted that the species in the genus Ostrinia are the most likely hosts. Of native North American species, Ostrinia obumbratalis is the only non-target species likely to be attacked by M. grandii. The predicted host range for North America matched the actual host range found in the field. This suggests that a careful literature review could be used as an important source of data on host range of parasitoid species proposed for introduction into a new environment.     

Assessment ofTrichogramma species for biological control of forest lepidopteran defoliators

In a laboratory study, we determined the potential of threeTrichogramma (Hymenoptera: Trichogrammatidae) species,T. brassicae Bezdenko,T. minutum Riley andT. nr.sibiricum Sorokina, for biological control against six species of forest lepidopteran pests, black army cutworm, hemlock looper, eastern spruce budworm, western spruce budworm, white-marked tussock moth, and gypsy moth. Females of each parasitoid species were offered eggs from each of the six host species. Parasitization and the effect of the host species on the emerging progeny were examined and recorded.Trichogramma minutum had the broadest host range and successfully parasitized four host species out of the six offered.Trichogramma nr.sibiricum had the narrowest host range and parasitized only two species of hosts. Of the six host species, black army cutworm was the most preferred by all threeTrichogramma species; white-marked tussock moth and gypsy moth were not parasitized by any parasitoids. There was a positive correlation between the size of female offspring and their corresponding egg complement in all three parasitoid species. The developmental time of parasitoids from egg to adult was influenced by both the parasitoid and host species. Our results suggest thatT. minutum has the greatest potential for biological control against various forest lepidopteran pests and that the black army cutworm may be the best target candidate for further study.     

A molecular phylogeny of the genus Lygodium (Schizaeaceae) with special reference to the biological control and host range testing of Lygodium microphyllum

Lygodium microphyllum, first naturalized in the 1960s, has aggressively invaded forest-dominated wetlands in southern and central Florida. The indeterminate growth of this invasive climbing fern creates thick rachis mats which climb over shrubs and trees smothering the underlying growth and carrying ground fires into the forest canopy. Foreign surveys for natural enemies identified 20 species of insects and two species of mites. Host range testing of three insect species and one mite included five Lygodium species (L. palmatum, L. volubile, L. cubense, L. venustum, and L. oligostachyum) native to the United States and the West Indies. A molecular phylogeny of the genus was conducted using the trnL intron and the trnL-F intergenic spacer of chloroplast DNA to determine the relationship of L. microphyllum to other Lygodium species. Three major clades appeared, one with L. palmatum and L. articulatum (the most basal), a second with L. reticulatum and L. microphyllum, and a third comprising the other species examined. Lygodium microphyllum appeared at the end of a long branch approximately equidistant from all species of interest preventing the correlation of genetic distance and host range behavior. However, inspection of the results of host range experiments showed a relationship between the presence of a related biotype or species of the insect on a related alternate host and the ability of that insect to develop on that alternate host. Ancestral host usage and ecological fitting are examined as possible explanations for the acceptance of L. palmatum by the tested musotimine insects.     

Larval performance of two leafroller species on known and potential hosts

Two univoltine leafroller species,Archips argyrospila Walker andArchips rosana L., were reared from eclosion to adulthood on known or potential host plants. Both species were able to complete development on eight of the ten plants tested, the exceptions being apricot and peach. Rates of development differed among hosts, and indicated a range of host suitability. Differences in female pupal weights were apparent following the different rearing treatments, but these did not correlate with subsequent female fecundity. When development was compared between the two leafroller species,A. rosana had higher survivorship, more rapid development and higher female fecundity than didA. argyrospila, regardless of host. These findings indicate thatA. rosana may successfully exploit a wider host range than doesA. argyrospila. Comparisons of survivorship and developmental rates among hosts for each species of leafroller suggest that both species may have the potential for host-range expansion.     

Responses of a generalist and a specialist parasitoid (Hymenoptera: Eucoilidae) to Drosophilid larval kairomones

Foraging parasitoids are thought to need more specific information than generalists on the presence, identity, availability, and suitability of their insect host species. In the present paper, we compare responses to host kairomones by two phylogenetically related parasitoid species that attack Drosophilidae and that differ in the width of their host range. As predicted, the behavioral response of the parasitoids to host kairomones reflected their difference in host range. The response of the specialist parasitoid Leptopilina boulardiwas restricted to contact kairomones from its natural hosts and one closely related species. In contrast, the generalist parasitoid Leptopilina heterotomaresponded to contact kairomones of a variety of Drosophilidae species.     

Morphological variation and cultural characteristics ofConiothyrium leucospermi associated with leaf spots of proteaceae

During an examination ofConiothyrium collections occurring on Proteaceae one species,C. leucospermi, was repeatedly encountered. However, it was not always possible to identify this species from host material alone, whereas cultural characteristics were found to be instrumental in its identification. Conidium wall ornamentation, which has earlier been accepted as crucial in species delimitation is shown to be variable on host material, making cultural comparisons essential. Using standard culture and incubation conditions,C. leucospermi is demonstrated to have a wide host range in the Proteaceae. In addition, microcyclic conidiation involving yeast-like budding from germinating conidia and hyphae in culture is newly reported for this species.     

Rapid evolution of host specificity in a parasitic nematode

Summary In eastern North America, the nematodeHowardula aoronymphium parasitizes four species of mushroom-breedingDrosophila:D. falleni andD. recens of the quinaria species group, andD. putrida andD. testacea of the testacea group. One strain ofH. aoronymphium, designated Mendon-87, was initially capable of infecting all four of these host species. After less than 3 years in laboratory culture usingD. falleni as the sole host, this strain had completely lost the ability to infectD. putrida. Two other nematode strains parasitizedD. falleni andD. putrida at equal rates. These results demonstrate the existence of genetic variation for host specificity within this nematode species. More importantly, they show that host specificity can evolve rapidly when only one host is available for parasitization. Ecological conditions are such that natural populations ofH. aoronymphium may comprise numerous host races, lineages incapable of parasitizing the full range of host species. However, I argue that such host races are probably ephemeral and thus unlikely to persist long enough to undergo speciation.     

Winter Host Plant Specialization in a Host-Alternating Aphid

The host alternating aphid, Rhopalosiphum padi (L.), feeds in summer on several different species of grasses but is monophagous on its winter host, Prunus padus L. The monophagy on P. padus could be a result of the restricted host range of the several different generations colonizing, or feeding, on this host during autumn–winter–spring. This study shows that the winter host plant specificity of R. padi is controlled mainly by the preference of the females remigrating (gynoparae) to the winter host, P. padus, in autumn. The other generations living on the winter host, i.e., sexual females, males, and spring generations, all accept a broader range of winter hosts. One alternative host plant, Prunus spinosa L., could be utilized by all generations associated with the winter host, except for the females remigrating at autumn.     

Relative host plant species use by the lantana biological control agent Aconophora compressa (Membracidae) across its native and introduced ranges

Aconophora compressa Walker (Hemiptera: Membracidae) was released in 1995 against the weed lantana in Australia, and is now found on multiple host plant species. The intensity and regularity at which A. compressa uses different host species was quantified in its introduced Australian range and also its native Mexican range. In Australia, host plants fell into three statistically defined categories, as indicated by the relative rates and intensities at which they were used in the field. Fiddlewood (Citharexylum spinosum L.: Verbenaceae) was used much more regularly and at higher densities than any other host sampled, and alone made up the first group. The second group, lantana (Lantana camara L.: Verbenaceae; pink variety) and geisha girl (Duranta erecta L.: Verbenaceae), were used less regularly and at much lower densities than fiddlewood. The third group, Sheena’s gold (another variety of D. erecta), jacaranda (Jacaranda mimosifolia D. Don: Bignoniaceae) and myoporum (Myoporum acuminatum R. Br.: Myoporaceae), were used infrequently and at even lower densities. In Mexico, the insect was found at relatively low densities on all hosts relative to those in Australia. Densities were highest on L. urticifolia, D. erecta and Tecoma stans (L.) Juss. ex Kunth (Bignoniaceae), which were used at similar rates to one another. It was found also on a few other verbenaceous and non-verbenaceous host species but at even lower densities. The relative rate at which Citharexylum spp. and L. urticifolia were used could not be assessed in Mexico because A. compressa was found on only one plant of each species in areas where these host species co-occurred. The low rate at which A. compressa occurred on fiddlewood in Mexico is likely to be an artefact of the short-term nature of the surveys or differences in the suites of Citharexylum and Lantana species available there. These results provide further incentive to insist on structured and quantified surveys of non-target host use in the native range of potential biological control agents prior to host testing studies in quarantine.     

Encapsulation and hemocyte numbers in three lepidopteran stemborers parasitized by Cotesia flavipes-complex endoparasitoids

Determination of the potential and actual host range of a natural enemy is crucial before its importation and release for biological control. We studied some of the factors that are important in determining the physiological host range of insect parasitoids attacking lepidopteran hosts. Our experimental system consisted of novel host-parasitoid associations, with two New World pyralid stalk borers, Diatraea saccharalis and D. grandiosella; one Old World crambid borer, Ostrinia nubilalis as hosts; and three Old World microgastrine braconids, Cotesia chilonis, C. sesamiae, and C. flavipes as parasitoids. Experiments on the chronology of encapsulation of the parasitoid progeny by host hemocytes indicated that lepidopteran stemborers that are taxonomically, behaviorally and ecologically very similar differ in their ability to encapsulate a parasitoid species. D. saccharalis encapsulated C. flavipes sometimes, whereas D. grandiosella consistently encapsulated C. sesamiae and C. flavipes. C. chilonis was not encapsulated by either Diatraea host. If encapsulation occurred it did not start until four days after parasitization and continued during the following days. O. nubilalis was an unsuitable host for all three parasitoid species; parasitoid eggs were killed within 24 hours of parasitization. O. nubilalis had nearly twice as many hemocytes present in the hemolymph compared to the Diatraea species. In many of the host-parasitoid combinations, there was an initial increase of hemocyte number soon after parasitization, which was not due to mechanical damage at oviposition. There was no correlation between total numbers of hemocytes present in the host hemolymph and the observed encapsulation levels. By understanding the encapsulation response we may be able to make better predictions about the host range of a parasitoid species before its release as a biological control agent.     

Puccinia kraussiana (Uredinales) onHeterosmilax gaudichaudiana,A new record of rust in Eastern Asia

Puccinia kraussiana, a rust fungus of Africa, was found in Taiwan for the first time in eastern Asia. It was parasiting on leaves ofHeterosmilax gaudichaudiana, on which all four stages were characteristically found. The host species,H. gaudichaudiana, newly added to the host range ofP. kraussiana, constitutes also a new entry to the phanerogamic flora of Taiwan.     

Suitability of lepidopteran stemborers for parasitization by novel-association endoparasitoids

Extensive study of insect immune systems has yielded abetter understanding of the mechanisms used by insects to defendagainst invaders. This knowledge can be used to predict hownatural enemies utilize potential hosts, which will aid in theplanning of biological control programs. Our experimental systemconsists of novel host-parasitoid associations, with two NewWorld pyralid stalk borers, Diatraea saccharalis and D.grandiosella; one Old World crambid borer, Ostrinia nubilalis;and three Old World microgastrine braconid parasitoids, Cotesiachilonis, C. sesamiae, and C. flavipes. Experiments on hostsuitability indicate that parasitoids that are taxonomically,behaviorally and ecologically very similar may differ in theirability to utilize a host of the same species. Likewise,utilization of related hosts can produce different outcomes for agiven parasitoid species. D. saccharalis is a suitable host forall three parasitoid species, whereas D. grandiosella oftenencapsulates C. sesamiae and C. flavipes. O. nubilalis is anunsuitable host for all three species. Different species ofparasitoids may use different factors at different times afterparasitization to counter the host's immune response. This studysuggests that the physiological host range of these parasitoids willbe narrow, thus limiting effects on non-target species. However,the lack of consistent patterns also shows that explicit testingwill be needed to determine host ranges.     

Parasitism by the mite Acarophenax lacunatus on beetle pests of stored products

Recent reports of the biocontrolpotential of the mite species Acarophenaxlacunatus (Cross & Krantz) (Prostigmata:Acarophenacidae) and the lack of biologicalstudies on this regulatory agent led to thepresent study carried out under laboratoryconditions. The objective of the investigationwas to assess the host range of A. lacunatus,so far only reported as egg parasite ofRhyzopertha dominica (F.) (Coleoptera:Bostrichidae). Four Coleoptera species ofstored cereals were used: R. dominica,Tribolium castaneum (Herbst) (Tenebrionidae),Cryptolestes ferrugineus (Stephens)(Laemophloeidae) and Oryzaephilus surinamensis (L.) (Cucujidae). The highest rates of eggparasitism were observed on R. dominica and T.castaneum, leading to a significant decrease ofpopulations of both species and reduced wheatweight loss. A. lacunatus was also able toparasitize eggs of C. ferrugineus, but not ofO. surinamensis. These results indicate abroader host range of A. lacunatus thaninitially suspected and also strengthen itspossibility of use in integrated pestmanagement programs in storage environments.     

Host location by larvae of the reptile tick Amblyomma vikirri (Acari: Ixodidae)

The newly described tick Amblyomma vikirri has a narrow host range, being found mainly on the Australian lizard, Egernia stokesii and rarely on another lizard, Tiliqua rugosa. Both hosts are in the family Scincidae. Larvae of A. vikirri were as successful in locating E. stokesii as T. rugosa from a range of release distances between 20 and 120 mm from the host. Over this range the proportion of ticks which successfully located hosts declined and the time taken by successful ticks to locate hosts increased with increasing release distance. From 60 mm, larvae of A. vikirri located four other lizard species from the families Scincidae and Agamidae and two non-living targets as successfully as they did E. stokesii. The only evidence that there was any host specificity in the searching behaviour of larvae of A. vikirri was that A. vikirri larvae spent less time paused and questing when they were searching for E. stokesii than when they were searching for T. rugosa. Aponomma hydrosauri, a tick which commonly infests T. rugosa but not E. stokesii, spent less time paused and questing when it was searching for T. rugosa than when it was searching for E. stokesii. However, the results overall suggest that the narrow host range of A. vikirri cannot be explained by any ability of the larvae of that species to discriminate between their natural host and other reptile species.     

Host plants of someRhinanthoideae (Scrophulariaceae) of eastern North America

Twenty three species in 11 genera were examined in the field to determine hosts. OnlyStriga asiatica andSeymeria cassioides have a narrow host range being restricted to grasses and pines, respectively. These are the only species which cause pronounced and sometimes serious host damage. The other species attach to a great diversity of hosts.     

The host range and selectivity of a parasitic plant: Rhinanthus minor L.

Summary Rhinanthus minor (Yellow-rattle) is a widespread hemiparasitic plant of grassland habitats throughout Britain. Association analysis of the dune vegetation at Holme-next-the-Sea in eastern England revealed only two potential host plants through positive association. In contrast direct examination of the root systems revealed haustorial connections with 20 host species. The number of species parasitized by one plant ranged from one to seven. Data from another four sites in Britain and one in central Europe indicate that the natural host range of R. minor encompasses at least 50 species from 18 families with 22% in the Leguminosae and 30% in the Gramineae. Comparison of the number of haustorial connections made to each species with the abundance of roots in the soil shows that R. minor is a highly selective parasite, but that the selectivity is not consistent between populations or between plants from different parts of the same population. The reasons for host selectivity are discussed.     

Experimental infections with the tropical monogenean,Gyrodactylus bullatarudis: Potential invader or experimental fluke?

Introduced exotic species have the potential to spread their associated parasites to native species which can be catastrophic if these hosts are immunologically naïve to the novel parasite. The guppy (Poecilia reticulata) has been disseminated worldwide outside of its native habitat and therefore could be an important source of infection to native fish species. Its parasite fauna is dominated by the ectoparasitic monogeneans, Gyrodactylus turnbulli and Gyrodactylus bullatarudis. The current study tested the host specificity of G. bullatarudis by experimentally infecting a range of isolated fish hosts, including temperate species. Surprisingly, the parasite was capable of establishing and reproducing, for several days, on the three-spined stickleback when transferred directly to this host. We also established that G. bullatarudis could be transmitted under aquarium conditions at both 25 °C and 15 °C. At the higher temperature, the parasite was even capable of reproducing on this atypical host. The implications of these findings are discussed in terms of host specificity, host switching and climate change.     

Influence of initial egg density and host size on the development of the gregarious parasitoid <Emphasis Type="Italic">Bracon hebetor</Emphasis> on three different host species

Bracon hebetor (Say) (Hymenoptera : Braconidae) is a gregarious parasitoid that attacks a variety of important lepidopterous pests of stored product and in the field. In this study the effect of host species, size and larval competition on parasitoid size, survival and development were investigated. In laboratory studies, wasp eggs at a range of densities, were placed on larvae of different weight of three Lepidoptera host species namely Adoxophyes orana (Fischer von Röslerstamm, Tortricidae), Plodia interpunctella (Hubner, Pyralidae) and, Lobesia botrana(Dennis & Schiffermueller, Tortricidae). On A. orana survival of immature parasitoids was very low at all densities and different host weights. On L. botrana survival progressively reduced as egg density increased at both host weights examined for this host. Survival on P. interpunctella was significantly affected by egg density but not by host weight. Initial egg density had a significant effect on the size of emerging adults from each rearing host. Smaller adult parasitoids emerged as egg density per larva increased. Larval host weight of P. interpunctella and A. orana had a significant effect on the size of emerging adult parasitoids mainly at the higher egg densities used in these experiments. The above results of host quality on fitness of parasitoid are discussed.     

Distribution, host specificity, and overwintering of Celatoria bosqi Blanchard (Diptera: Tachinidae), a South American parasitoid of Diabrotica spp. (Coleoptera: Chrysomelidae: Galerucinae)

The genus Diabrotica (Coleoptera: Chrysomelidae) includes a great number of pest species, including some of the most important crops pests of the Americas. However, only five parasitoid species have been recorded for it. The parasitoid Celatoria bosqi Blanchard was the first parasitoid described from Diabrotica spp. in South America, where substantial parasitism has been observed. C. bosqi has been collected almost throughout the South American distribution of its main host, Diabrotica speciosa (Germar), in an area that includes temperate and tropical lowlands, and semiarid to humid highlands. Three Diabrotica species were found to host the parasitoid, D. speciosa (Germar), Hystiopsis sp., and Diabrotica viridula (F.), with a total parasitism of 2.60, 5.55, and <0.02%, respectively. Laboratory experiments with field beetles and puparia, reared in the laboratory, indicate that C. bosqi overwinters obligatorily in overwintering adult host beetles, remaining quiescent in its live host below developmental temperatures. Based on the known climatic range of C. bosqi, and its requirement of adult overwintering hosts, a potential distribution in North America is projected.