Host plant use and possibly defensive horns in <i>Trichochermes magna</i>(Psilidae: Homoptera)

ABSTRACT. Tent caterpillars are generalists across their full host range, but display local host plant preferences. We present evidence for a new host plant record, wax currant (Ribes cereum), for western tent caterpillars (Malacosoma californicum) along the Colorado Front Range. We tested the suitability of wax currant as a host plant for western tent caterpillars as compared to chokecherry (Prunus virginiana), an abundant and commonly used host plant. We measured the density of tent caterpillar tents in areas where both host plants occur to assess host plant use. We reared tent caterpillar larvae on both host plants and measured fitness effects due to host plant quality (survival, pupal mass) and natural enemies (parasitism). We did not find a relationship between host plant abundance and use by tent caterpillars and found no evidence for a preference for either host plant. We found that western tent caterpillars do not differ in pupal mass when reared on chokecherry and on wax currant in a laboratory s...

While the host plant use of insect herbivores is important for understanding their interactions and coevolution, field evidence of these preferences is limited for generalist species. Molecular diet analysis provides an effective option for gaining such information, but data from field‐sampled individuals are often greatly affected by the local composition of their host plants. The polyphagous mirid bug Apolygus lucorum (Meyer‐Dür) seasonally migrates across the Bohai Sea, and molecular analysis of migrant bugs collected on crop‐free islands can be used to estimate the host plant use of A. lucorum across the large area (northern China) from where these individuals come. In this study, the host plant use of A. lucorum adults was determined by identifying plant DNA using a three‐locus DNA barcode (rbcL, trnH‐psbA, and ITS) in the gut of migrant individuals collected on Beihuang Island. We successfully identified the host plant families of A. lucorum adults, and the results indicated that captured bugs fed on at least 17 plant families. In addition, gut analyses revealed that 35.9% of A. lucorum individuals fed on multiple host plants but that most individuals (64.1%) fed on only one plant species. Cotton, Gossypium hirsutum L., DNA was found in 35.8% of the A. lucorum bugs examined, which was much higher than the percentage of bugs in which other host plants were found. Our work provides a new understanding of multiple host plant use by A. lucorum under natural conditions, and these findings are available for developing effective management strategies against this polyphagous pest species.

Population divergence can occur due to mechanisms associated with geographic isolation and/or due to selection associated with different ecological niches. Much of the evidence for selection-driven speciation has come from studies of specialist insect herbivores that use different host plant species; however, the influence of host plant use on population divergence of generalist herbivores remains poorly understood. We tested how diet breadth, host plant species and geographic distance influence population divergence of the fall webworm (Hyphantria cunea; FW). FW is a broadly distributed, extreme generalist herbivore consisting of two morphotypes that have been argued to represent two different species: black-headed and red-headed. We characterized the differentiation of FW populations at two geographic scales. We first analysed the influence of host plant and geographic distance on genetic divergence across a broad continental scale for both colour types. We further analysed the influence of host plant, diet breadth and geographic distance on divergence at a finer geographic scale focusing on red-headed FW in Colorado. We found clear genetic and morphological distinction between red- and black-headed FW, and Colorado FW formed a genetic cluster distinct from other locations. Although both geographic distance and host plant use were correlated with genetic distance, geographic distance accounted for up to 3× more variation in genetic distance than did host plant use. As a rare study investigating the genetic structure of a widespread generalist herbivore over a broad geographic range (up to 3,000km), our study supports a strong role for geographic isolation in divergence in this system.

Studies on strategies of host plant use in sympatric-related species are significant to the theory of sympatric speciation. Altica fragariae Nakane and Altica koreana Ogloblin are sympatric closely related flea beetles found in Beijing, northern China. All their recorded host plants are in the subfamily Rosoideae of the Rosaceae, so we regard them as a model system to study interactions between herbivorous insects and plant-insect co-evolution. We conducted a set of experiments on the host preference and performance of these flea beetles to study whether these closely related species have the ability to use sympatric novel host plants and whether monophagous and oligophagous flea beetles use the same strategy in host plant use. Oviposition preference experiments showed that A. koreana, a monophagous flea beetle, displayed high host fidelity. However, A. fragariae, which is oligophagous, often made "oviposition mistakes," ovipositing on nonhost plants such as Potentilla chinensis, the host plant of A. koreana, although normal host plants were preferred over novel ones. Larval performance studies suggested that A. fragariae was able to develop successfully on P. chinensis. Feeding experiences of larvae had no effect on feeding preference, oviposition preference, and fecundity of adults. However, females were impaired in their reproductive ability when fed on nonhost plants. Therefore, A. fragariae finished their development of larval stages on P. chinensis and came back to their primary host plant, Duchesnea indica, for feeding and reproduction after eclosion.

Abstract.1. The butterflyMelitaea cinxiauses two host plant species in the Åland Islands of south‐west Finland. Survey data show that host plant use is spatially variable and that the two species are not used (fed on byM.cinxialarvae) in proportion to their abundances. The pattern of host plant use byM.cinxiahas been attributed in part to plant distribution and spatial variation of butterfly oviposition preference.2. The additional roles that may be played by spatial variation in host plant quality and larval physiology (performance ability) were investigated. Seven years of field survey data and a series of laboratory experiments were used to demonstrate that neither of these variables contributed measurably to the observed pattern of host plant use.3. Specifically, while there was great variation among individuals in both the performance ability of caterpillars and host plant suitability, the two plant species appeared equally suitable, there was no within‐species geographic variation in plant suitability, and there was no evidence for adaptation of caterpillars to the locally used host plants.

There is considerable interspecific variation in larval host plants (Passifloraceae) used among Heliconius erato (Lepidoptera: Nymphalidae) populations. This study evaluates the role of learning and the influence of interspecific variation in host plant attributes on such local specialization in H. erato host plant choices. Experiments were conducted under laboratory, insectary, and field conditions, with the two most widely used host plants in southern Brazil (Passiflora suberosa Linnaeus and Passiflora misera Humbold, Bonpland et Kunth). Larval feeding preference and induction were evaluated through choice tests for all instars. Oviposition was evaluated in relation to host plant preference, Hopkins host-selection principle, and conditioning time (for 3, 7, 11, or 15 days). Also, oviposition choice was tested regarding density, intemode length, and presence of intact terminal bud on P. suberosa and P. misera shoots. Both larvae and adults of H. erato phyllis showed preference for P. misera compared to P. suberosa, under all conditions. Larval feeding preference could not be induced for most instars. The Hopkins’ effect was not detected and oviposition choice could not be conditioned. Females alternated use of host plant species as a function of variation in either density or presence of terminal buds on shoots. Thus, our data indicate host plant preference in H. erato phyllis is not learned but innate. Therefore, we concluded that variation in local use of host plant by this butterfly in southern Brazil results from qualitative and quantitative variation of the passion vine species.

The genus Eustigmaeus Berlese, 1910 represents the unique phytophagous group within the superfamily Raphignathoidea. Four species within this genus have been known to inhabit mosses and feed on them as larvae, nymphs, and adults. However, the interactions with mosses have remained poorly understood. In order to reveal the diversity and host-plant use of the moss-feeding species, we conducted an extensive field study in Japan. This study revealed an array of moss-feeding species inhabiting various moss species, with 10 morphologically distinctive species newly documented in Japan. Through DNA barcoding based on cytochrome c oxidase subunit I (COI) sequences, these morphospecies were recovered as distinct entities. Notably, the host-plant use of four species was elucidated. Among these, Eustigmaeus sp. 9 exhibited polyphagy, while three species (Eustigmaeus spp. 1-3) demonstrated varying degrees of host specificity, each using moss species from the Hypnales, Philonotis, and Dicranidae, respectively. While a few moss-feeding species were frequently found in the same geographic area, more than one species rarely co-occurred within the same moss colonies. Eustigmaeus offers a unique study system, with its diverse moss-feeding species and indications of specific host plant use. Consequently, the moss-feeding Eustigmaeus serves as a valuable model for exploring the macroevolutionary patterns underlying diversification in moss-feeding arthropods.

The view of (insect) populations as assemblages of local subpopulations connected by gene flow is gaining ground. In such structured populations, local adaptation may occur. In phytophagous insects, one way in which local adaptation has been demonstrated is by performing reciprocal transplant experiments where performance of insects on native and novel host plants are compared. Trade‐offs are assumed to be responsible for a negative correlation in performance on alternative host plants. Due to mixed results of these experiments, the importance of trade‐offs in host plant use of phytophagous insects has been under discussion. Here we propose that another genetic mechanism, the evolution of coadapted gene complexes, might also be associated with local adaptation. In this case, however, transplant experiments might not reveal any local adaptation until hybridization takes place. We review the results we have obtained in our work on the host plant use of the flea beetle Phyllotreta nemorum L. (Coleoptera: Chrysomelidae: Alticinae), and propose a hypothesis involving coadapted genes to explain the distribution of genes that render P. nemorum resistant to defences of one of its host plants, Barbarea vulgaris R. Br. (Cruciferae).

Dose-dependent dynamics of densovirus infection in two nymphalid butterfly species utilizing native or exotic host plants

Host plant use and availability were determined in early nymphal and adult-stage Schistocerca emarginata (=lineata) (Orthoptera: Acrididae) populations at six localities in Texas, USA. Early instar nymphal populations were feeding almost exclusively on either Ptelea trifoliata (Rutaceae) or Rubus trivialis (Rosaceae). This study represents the first demonstration of a geographic structure of host plant specificity in a polyphagous grasshopper. Recognizing this geographic structure required investigations of both developmental and geographical variation in host plant use. Nymphal diet breadths were significantly less than adult diet breadths at four of six localities and smaller overall when pooled nymphal and adult diet breadths were compared among sites. Neither restricted nymphal mobility nor host plant availability accounted for the observed differences in host plant use between developmental stages and among localities. Evidence suggests that the differences in host use among populations are due to host-plant-associated genetic differentiation.

Insect herbivores that are tended by ants exhibit a range in host plant use from specialists to extreme generalists. Potential factors that may influence relative suitability of different host plants include the presence or absence of ants and the size of aggregations formed by nymphs feeding on a host. We used a membracid-ant mutualism to test whether presence of ants or aggregation size (from 20 to 320 nymphs per plant) resulted in variable selection for host use by the generalist membracid Publilia modesta (Uhler) (Hemiptera: Membracidae) that feeds on host species Chrysothamnus viscidiflorus (Hook.) Nutt. and Wyethia spp., among others. Survival of nymphs as mid-instars and the percentage of nymphs developing into adults on C. viscidiflorus were greater for nymphs in small than in large aggregations. Survival of nymphs as mid-instars on Wyethia spp. was also greater in smaller aggregations, but the percentage of nymphs developing into adults was independent of aggregation size. The presence of the tending ant, Formica obscuripes Forel, had no effect on the percentage of nymphs developing into adults during this experiment. The survival of membracids on C. viscidiflorus was always greater than survival of membracids on Wyethia spp., regardless of aggregation size or presence of ants; the favored host did not vary in a way that would explain the observed generalized host plant use. C. viscidiflorus was a superior host plant, because it senesced after nymphs became adults, whereas Wyethia spp. senesced early enough that most nymphs died before reaching adulthood.

Potential host plants of the polyphagous lepidopteran Helicoverpa punctigera (Wallengren) were surveyed in two ways. A broad survey, conducted in southern Queensland and northern New South Wales, indicated that H. punctigera larvae were present on relatively few plant species. A detailed survey of host plant use in a non‐cropping area in which H. punctigera was numerous demonstrated restricted host plant use by this species. The density of H. punctigera on its principal host in the area, the indigenous daisy Ixiolaena brevicompta F. Muell., was much higher (as measured per unit of time searched) than on other plant species available. Also, I. brevicompta was used regularly by H. punctigera after rainfall events. Ixiolaena brevicompta represents a new host record and on the basis of the pattern of its use by H. punctigera should be considered a ‘primary host plant’ of this noctuid. In cropping areas sampled, usually more than one plant species hosted H. punctigera regularly and in large numbers. Usually a crop species was included (e.g. cotton and chick pea). Alternative hosts in cropping areas were Sonchus oleraceus L. (sowthistle) and possibly the native legume Sesbania cannabina (Retz.) Poiret. Our results imply that the polyphagy of H. punctigera is probably not as extensive as previously claimed. The criteria for inclusion of a plant species as a primary host for H. punctigera need to include the regularity of use of that species and the relative abundance of eggs and larvae on it. We suggest that an understanding of the host‐searching mechanism of this species will be best achieved through study of the interaction of H. punctigera with its indigenous primary hosts. The surveys also yielded information on host plants of two other heliothine noctuids, H. armigera (Hübner) and Australothis rubrescens (Walker), and this is also presented.

Leaf beetles in the genus Phratora differ in host plant use and in the chemical composition of their larval defensive secretion. Most species specialize on either poplars or willows (family Salicaceae), but two species feed on birch (family Betulaceae). Phratora vitellinae utilizes salicylates from the host plant to produce its larval secretion, which contains salicylaldehyde, while other Phratora species produce an autogenous secretion. To reconstruct the evolutionary history of host plant use and the larval secretion chemistry in this genus, we sequenced 1383 base pairs of the mt cytochrome oxidase I gene for six European and one North American Phratora species and three outgroup taxa. Bootstrap values of the complete nucleotide sequence were 99-100% for six of eight nodes in the maximum parsimony tree. They were 71% and 77% for the two other nodes. The maximum parsimony tree and the maximum likelihood tree based on nucleotide sequence showed the same relationships as a maximum parsimony tree based on the amino acid sequence. Beetle phylogeny overlapped broadly with host plant taxonomy and chemistry, and it revealed historical constraints influencing host plant use. However, there was one host shift from the willow family (Salicaceae) to the birch family (Betulaceae). The use of host plant phenol glycosides for the larval defensive secretion evolved along the lineage that led to P. vitellinae. Phratora vitellinae feeds on the taxonomically widest range of host plants, which are characterized by moderate to high levels of salicylates. The results support the hypothesis that the use of salicylates for the larval secretion evolved twice independently in chrysomeline leaf beetles.

There is a great number of Charaxinae (Lepidoptera: Nymphalidae) species in the tropics whose larvae feed on several plant families. However the genus Anaea is almost always associated with Croton species (Euphorbiaceae). This work describes patterns of host plant use by immature and adult abundance on different vertical strata of sympatric Anaea species in a forest of Southeastern Brazil. Quantitative samples of leaves were taken in April/1999 and May/2000 to collect eggs and larvae of four Anaea species on C. alchorneicarpus, C. floribundus and C. salutaris in a semideciduous forest. Sampled leaves were divided into three classes of plant phenological stage: saplings, shrubs and trees. The results showed that the butterfly species are segregating in host plant use on two scales: host plant species and plant phenological stages. C. alchorneicarpus was used by only one Anaea species, whereas C. floribundus was used by three species and C. salutaris by four Anaea species. There was one Anaea species concentrated on sapling, another on sapling/shrub and two others on shrub/tree leaves. Adults of Anaea were more frequent at canopy traps but there were no differences among species caught in traps at different vertical positions. This work supplements early studies on host plant use among Charaxinae species and it describes how a guild of closely related butterfly species may be organized in a complex tropical habitat.

Herbivore diet breadth has been hypothesised to be driven by plant quality (physiological efficiency hypothesis) or natural enemies (enemy‐free space hypothesis). These hypotheses on their own are now known to be insufficient explanations for the range of herbivore diet breadths and more integrative approaches consider how trade‐offs and ecological contingencies shape host plant use by herbivores. I carried out laboratory experiments to test whether physiological efficiency and defence against natural enemies trade off when larvae of the ornate bella moth,Utetheisa ornatrix(L.) (Lepidoptera: Erebidae), a genus‐specialist pod‐boring herbivore, feeds on leaves and pods of fourCrotalariaspecies (Fabaceae). Different components ofU.ornatrixphysiological efficiency traded off amongst each other and with defence against adult predation by the spiderLycosa erythrognathaLucas (Araneae: Lycosidae).Crotalaria micansLink provided the highest survival and fast development but low pupal weight and defence.Crotalaria ochroleucaG. Don provided poor survival, slow development, and low defence, but high pupal weight.Crotalaria pallidaAiton provided slow development and intermediate survival and pupal weight, but the best defence.Crotalaria vitellinaKer Gawl. provided high survival, fast development, and efficient defence, but low pupal weight. Compared to leaves, feeding on green seeds provided greater defence against predation, and developmental benefits, independent of host plant species, but there was a large cost in boring into the pod to reach the seeds. I therefore provide evidence that the trade‐offs among physiological efficiency components and between physiological efficiency and defence could maintain multiple host plant use in specialist herbivores. I also show that feeding on green seeds incurs a cost through pod‐boring but gives the moth benefits in adult defence. Therefore, the evolution of host plant use and pod‐boring in insect herbivores could be driven by defence as much as by physiological efficiency.