Experimental assemblage of novel plant–herbivore interactions: ecological host shifts after 40 million years of isolation

Geographic isolation is the first step in insect herbivore diet specialization. Such specialization is postulated to increase insect fitness, but may simultaneously reduce insect ability to colonize novel hosts. During the Paleocene‐Eocene, plants from the order Zingiberales became isolated either in the Paleotropics or in the Neotropics. During the Cretaceous, rolled‐leaf beetles diversified in the Neotropics concurrently with Neotropical Zingiberales. Using a community of Costa Rican rolled‐leaf beetles and their Zingiberales host plants as study system, we explored if previous geographic isolation precludes insects to expand their diets to exotic hosts. We recorded interactions between rolled‐leaf beetles and native Zingiberales by combining DNA barcodes and field records for 7450 beetles feeding on 3202 host plants. To determine phylogenetic patterns of diet expansions, we established 20 experimental plots in the field, in which we planted plots five exotic Zingiberales, recording beetles feeding on these exotic hosts. In the laboratory, using both native and exotic host plants, we reared a subset of insect species that had expanded their diets to the exotic plants. The original plant–herbivore community comprised 24 beetle species feeding on 35 native hosts, representing 103 plant–herbivore interactions. After exotic host plant introduction, 20 percent of the beetle species expanded their diets to exotic Zingiberales. Insects only established on exotic hosts that belong to the same plant family as their native hosts. Laboratory experiments show that beetles are able to complete development on these novel hosts. In conclusion, rolled‐leaf beetles are preadapted to expand their diets to novel host plants even after millions of years of geographic isolation.     

Performance of Danaini larvae is affected by both exotic host plants and abiotic conditions

The consequences of the introduction of invasive plants for the diet of herbivorous insects have been little explored in nature where, potentially, abiotic and biotic factors operate. In this study, we examined the larval performance of two Neotropical Danaini butterflies when using either a native or an exotic Apocynaceae species as host plant in both field and laboratory experiments. Hosts greatly differ in their amount of latex exudation and other physicochemical traits, as well as in the amount of evolutionary time they have interacted with herbivores. First, herbivore performance on the hosts was investigated under laboratory conditions. Larvae of both Danaini species took more time to develop on the exotic host; larval survivorship did not vary between hosts. Second, first instar survivorship on both hosts was evaluated in two field sites, one site per host. To do so, in both sites half of the larvae were bagged (protected against both abiotic and biotic factors) while the remainder were nonbagged (exposed). The interaction between larval exposure with the use of the exotic host reduced larval survival. We concluded that the combined effects of host plant traits and abiotic factors reduced survival of herbivores in field conditions. Therefore, the performance of herbivores when using hosts of different origins should be considered together with the multiple ecological factors found in natural environments, as these factors can modify the result of plant–herbivore interactions.     

The population ecology of novel plant–herbivore interactions

Many native herbivores are known to attack exotic plants, and we can expect these interactions to occur with increasing frequency in coming years as invasive plants become naturalized and new invaders arrive in native communities. In some cases, evolutionary biologists and ecologists have learned a great deal from insects adapting to novel hosts. However, there is more to be learned and we suggest that the ecological study of exotic host colonization by native insects has been impeded by a lack of focus in the questions being asked, and also from overlap with other areas of plant–insect ecology, including the study of specialization. In the present paper, a conceptual model is described for the colonization of a novel host‐plant, which focuses on the relationship between occupancy and availability. Occupancy is the fraction of patches of novel hosts that are utilized by an herbivore, and availability is the abundance or presence of a novel host on the landscape. Considering the slope of that relationship (between occupancy and availability), hypotheses are suggested that involve dispersal and, most important, population growth rate of an insect herbivore associated with an exotic host. A focus on the occupancy–availability relationship highlights the strengths and weaknesses of common experimental approaches, such as preference–performance experiments. Suggestions for future work are offered that include integration with evolutionary theory and exploration of more complex demographic and ecological scenarios.     

Future danger posed by fungi in the Ophiostomatales when encountering new hosts

The Ophiostomatales contain pathogens that threaten forests world-wide. Global trade increases encounters with new hosts, with potential devastating consequences. We assessed the danger posed by the movement of Ophiostomatales between different host trees in South Africa. We tested the pathogenicity of five fungal species from native South African trees, and three from exotic trees, on various native and exotic trees. To evaluate the potential of fungi to move to new hosts, we investigated the strength of their associations with arthropod vectors. Results indicate that many fungal species are pathogens of newly encountered and distantly-related hosts. Encounters of pathogens with new hosts are less likely when host plants are distantly related, and outside the host range of boring beetle vectors, which also reduces the chances of vectoring by phoretic mite associates. However, pathogens associated with numerous mite species and wounds are more likely to encounter new hosts and pose future threats.     

Nonadaptive radiation: Pervasive diet specialization by drift in scale insects?

At least half of metazoan species are herbivorous insects. Why are they so diverse? Most herbivorous insects feed on few plant species, and adaptive host specialization is often invoked to explain their diversification. Nevertheless, it is possible that the narrow host ranges of many herbivorous insects are nonadaptive. Here, we test predictions of this hypothesis with comparative phylogenetic analyses of scale insects, a group for which there appear to be few host‐use trade‐offs that would select against polyphagy, and for which passive wind‐dispersal should make host specificity costly. We infer a strong positive relationship between host range and diversification rate, and a marked asymmetry in cladogenetic changes in diet breadth. These results are consonant with a system of pervasive nonadaptive host specialization in which small, drift‐ and extinction‐prone populations are frequently isolated from persistent and polyphagous source populations. They also contrast with the negative relationship between diet breadth and taxonomic diversification that has been estimated in butterflies, a disparity that likely stems from differences in the average costs and benefits of host specificity and generalism in scale insects versus butterflies. Our results indicate the potential for nonadaptive processes to be important to diet‐breadth evolution and taxonomic diversification across herbivorous insects.     

Host plant adaptation during contemporary range expansion in the monarch butterfly

Herbivores that have recently expanded their host plant ranges provide opportunities to test hypotheses about the evolution of host plant specialization. Here, we take advantage of the contemporary global range expansion of the monarch butterfly (Danaus plexippus) and conduct a reciprocal rearing experiment involving monarch populations with divergent host plant assemblages. Specifically, we ask the following questions: (1) Do geographically disparate populations of monarch butterflies show evidence for local adaptation to their host plants? If so, what processes contribute to this pattern? (2) How is dietary breadth related to performance across multiple host species in monarch populations? (3) Does the coefficient of variation in performance vary across sympatric versus allopatric hosts? We find evidence for local adaptation in larval growth rate and survival based on sympatric/allopatric contrasts. Migratory North American monarchs, which have comparatively broad host breadth, have higher mean performance than derived nonmigratory populations across all host plant species. Monarchs reared on their sympatric host plants show lower coefficient of variation in performance than monarchs reared on allopatric hosts. We focus our discussion on possible mechanisms contributing to local adaptation to novel host plants and potential explanations for the reduction in performance that we observed in derived monarch populations.     

Variation of butterfly diet breadth in relation to host-plant predictability: results from two faunas

Host‐plant data for North American and Australian butterflies were used to test the hypothesis that larval diet breadth increases with decreasing resource predictability (where the latter was estimated by host‐plant growth‐form/duration). For each region in turn we compared the diet breadths of butterflies which utilise herbaceous host‐plants with those of species having woody hosts. For North America alone we also compared the diet breadths of species having annual hosts with those utilising perennial hosts, and the diets of species having herbaceous‐annual hosts with those using woody‐perennial hosts. Studies of diet breadth may be biased by the host taxonomic level which contributes most to the diet index used. For example, the results of analyses which employ indices based on numbers of families of hosts utilised may differ from those using indices based on counts of host species or genera. To investigate this potential problem we performed cross‐species analyses where diet breadth was defined in turn as the number of host species, genera, or families eaten. We found that using different taxonomic levels did give inconsistent results. To avoid this we employed phylogenetic diet breadth indices in comparative analyses of Independent Contrasts. The former incorporate information from the whole of the host‐plant phylogeny, whilst the comparative method eliminates any confounding effects of butterfly phylogeny. The results indicated that there is a phylogenetic component to butterfly diet breadth. They also largely differed from those of the cross‐species investigations, although there were similarities (i.e. results differed between regions and varied according to whether the whole fauna or just endemics were investigated). Our results suggested that in both regions, non‐endemics which feed on herbaceous plants have wider diet breadths than non‐endemics which utilise woody hosts. However, we found no consistent evidence that the diet breadths of endemics increase with decreasing resource predictability (as estimated here).     

The relationship between diet breadth and geographic range size in the butterfly subfamily Nymphalinae--a study of global scale

The "oscillation hypothesis" has been proposed as a general explanation for the exceptional diversification of herbivorous insect species. The hypothesis states that speciation rates are elevated through repeated correlated changes--oscillations--in degree of host plant specificity and geographic range. The aim of this study is to test one of the predictions from the oscillation hypothesis: a positive correlation between diet breadth (number of host plants used) and geographic range size, using the globally distributed butterfly subfamily Nymphalinae. Data on diet breadth and global geographic range were collected for 182 Nymphalinae butterflies species and the size of the geographic range was measured using a GIS. We tested both diet breadth and geographic range size for phylogenetic signal to see if species are independent of each other with respect to these characters. As this test gave inconclusive results, data was analysed both using cross-species comparisons and taking phylogeny into account using generalised estimating equations as applied in the APE package in R. Irrespective of which method was used, we found a significant positive correlation between diet breadth and geographic range size. These results are consistent for two different measures of diet breadth and removal of outliers. We conclude that the global range sizes of Nymphalinae butterflies are correlated to diet breadth. That is, butterflies that feed on a large number of host plants tend to have larger geographic ranges than do butterflies that feed on fewer plants. These results lend support for an important step in the oscillation hypothesis of plant-driven diversification, in that it can provide the necessary fuel for future population fragmentation and speciation.     

The evolution of novel host use is unlikely to be constrained by trade‐offs or a lack of genetic variation

The genetic and ecological factors that shape the evolution of animal diets remain poorly understood. For herbivorous insects, the expectation has been that trade‐offs exist, such that adaptation to one host plant reduces performance on other potential hosts. We investigated the genetic architecture of alternative host use by rearing individual Lycaeides melissa butterflies from two wild populations in a crossed design on two hosts (one native and one introduced) and analysing the genetic basis of differences in performance using genomic approaches. Survival during the experiment was highest when butterfly larvae were reared on their natal host plant, consistent with local adaptation. However, cross‐host correlations in performance among families (within populations) were not different from zero. We found that L. melissa populations possess genetic variation for larval performance and variation in performance had a polygenic basis. We documented very few genetic variants with trade‐offs that would inherently constrain diet breadth by preventing the optimization of performance across hosts. Instead, most genetic variants that affected performance on one host had little to no effect on the other host. In total, these results suggest that genetic trade‐offs are not the primary cause of dietary specialization in L. melissa butterflies.     

Potential novel hosts for the lily leaf beetle Lilioceris lilii Scopoli (Coleoptera: Chrysomelidae) in eastern North America

Abstract.  1. Introduced insects often incorporate native plants into their diets and might be expected to show a predilection for novel hosts that are phylogenetically related to their normal hosts. The lily leaf beetle, Lilioceris lilii (Coleoptera: Chrysomelidae), is an introduced pest of cultivated lilies. Oviposition behaviour, larval behaviour, and development of L. lilii was examined on a range of potential host plants, as well as on the normal host, Asiatic hybrid lilies Lilium sp.
2. Neonate larval feeding behaviour was quantified on 15 food plant species: 10 from the Liliales, three from the Asparagales and two eudicots. Larvae fed plants closely related to the genus Lilium were more likely to initiate feeding, less likely to abandon their food leaf, and consumed more leaf area.
3. In no-choice tests, females oviposited on the novel hosts Lilium philadelphicum , Medeola virginiana , Clintonia borealis , Streptopus amplexifolius , and Polygonatum biflorum ; however, all but L. philadelphicum received very few eggs. Non- Lilium novel hosts were not used for oviposition when presented along with Asiatic lilies in choice tests.
4. A single individual was reared to the adult stage on the novel host S. amplexifolius . Several larvae survived to the pupal stage on M. virginiana , although no adults emerged from those pupae. Larvae reared on the native wood lily L. philadelphicum performed equally well or better than on the Asiatic cultivar.
5. Our results indicate that the lily leaf beetle poses a threat to native Liliaceae. Several native Lilium species, including L. philadelphicum , are threatened or endangered in certain jurisdictions throughout their range; these species should be monitored closely for colonisation by the beetle.     

Insects as alternative hosts for phytopathogenic bacteria

Phytopathogens have evolved specialized pathogenicity determinants that enable them to colonize their specific plant hosts and cause disease, but their intimate associations with plants also predispose them to frequent encounters with herbivorous insects, providing these phytopathogens with ample opportunity to colonize and eventually evolve alternative associations with insects. Decades of research have revealed that these associations have resulted in the formation of bacterial-vector relationships, in which the insect mediates dissemination of the plant pathogen. Emerging research, however, has highlighted the ability of plant pathogenic bacteria to use insects as alternative hosts, exploiting them as they would their primary plant host. The identification of specific bacterial genetic determinants that mediate the interaction between bacterium and insect suggests that these interactions are not incidental, but have likely arisen following the repeated association of microorganisms with particular insects over evolutionary time. This review will address the biology and ecology of phytopathogenic bacteria that interact with insects, including the traditional role of insects as vectors, as well as the newly emerging paradigm of insects serving as alternative primary hosts. Also discussed is one case where an insect serves as both host and vector, which may represent a transitionary stage in the evolution of insect-phytopathogen associations.     

Host range evolution is not driven by the optimization of larval performance: the case of <Emphasis Type="Italic">Lycaeides melissa</Emphasis> (Lepidoptera: Lycaenidae) and the colonization of alfalfa

Herbivorous insects that have recently incorporated novel hosts into their diet provide unique opportunities for understanding factors that promote or constrain the evolution of niche breadth. Lycaeides melissa has colonized both cultivated and feral alfalfa (Medicago sativa) throughout much of North America within the past 200 years. We investigated the quality of the novel host as a resource for juvenile development, and asked if the novel host is a preferred host for oviposition relative to a native host (Astragalus canadensis). Larval-performance and oviposition-preference were examined using L. melissa individuals from a population associated with both M. sativa and A. canadensis, and oviposition-preference was also examined in another population associated exclusively with M. sativa. In addition, we investigated the effects of M. sativa and A. canadensis flowers on both preference and performance. Only one of the hosts, M. sativa, has flowers that are accessible to nectaring butterflies, and we hypothesized that the presence of flowers could affect female behavior. We find that the novel host is a relatively poor larval resource: adults that were reared as larvae on M. sativa were roughly one-third the size of adults that were reared on the native host, A. canadensis. The native host, Astragalus canadensis, is the preferred host in choice experiments involving only foliage. However, when flowers were included in preference assays, the native and novel hosts received similar numbers of eggs. Thus, the presence of flowers on hosts in the field might influence the utilization of a novel and inferior larval resource. These results are consistent with a model in which host shifts are driven by adult behavior that does not directly optimize larval performance.     

视觉信号在昆虫检测植物寄主中的作用

植物为数十万种昆虫提供各种资源,如食物、交配、产卵和躲避天敌的场所。目前对昆虫检测植物寄主的研究主要关注昆虫嗅觉系统和植物寄主挥发物之间的相互作用,对昆虫视觉系统发挥的作用关注较少。近年来,对昆虫视觉器官、光行为反应及分子生物学的研究表明,昆虫具有优异的视觉能力,能够辨别植物寄主的颜色、大小和轮廓,应该将视觉纳入昆虫检测植物寄主的研究中。昆虫能够利用视觉信号准确检测寄主,远距离时,主要依靠植物寄主轮廓检测寄主,近距离时,寄主的大小、颜色和形状发挥重要作用。利用昆虫视觉识别寄主的专一性研制诱捕装置,可为害虫的监测和防治提供一定的理论基础。     

Predicting novel trophic interactions in a non‐native world

Humans are altering the global distributional ranges of plants, while their co‐evolved herbivores are frequently left behind. Native herbivores often colonise non‐native plants, potentially reducing invasion success or causing economic loss to introduced agricultural crops. We developed a predictive model to forecast novel interactions and verified it with a data set containing hundreds of observed novel plant–insect interactions. Using a food network of 900 native European butterfly and moth species and 1944 native plants, we built an herbivore host‐use model. By extrapolating host use from the native herbivore–plant food network, we accurately forecasted the observed novel use of 459 non‐native plant species by native herbivores. Patterns that governed herbivore host breadth on co‐evolved native plants were equally important in determining non‐native hosts. Our results make the forecasting of novel herbivore communities feasible in order to better understand the fate and impact of introduced plants.     

Fruit‐breeding drosophilids (Diptera) in the Neotropics: playing the field and specialising in generalism?

1. Species of Drosophilidae are frequently used as model organisms, but their relationships with the environment, particularly in immature stages, remain poorly known. 2. This is the most comprehensive survey to date of fruit‐breeding drosophilids and their hosts in the Neotropics. Drosophilid host‐utilisation patterns were analysed as to geographic origin (native versus exotic) and level of specialisation. 3. The 180 species of plants recorded as drosophilid hosts are distributed across the main Angiosperm lineages and fleshy‐fruited orders; plant families that hosted the greatest number of drosophilid species were Arecaceae, Moraceae, and Myrtaceae. The 100 nominal drosophilid species recorded breeding in fruits belong to just over one‐third of Neotropical genera; most species (91) belong to Drosophila. Drosophilid species with the greatest resource breadth were Drosophila simulans, Drosophila nebulosa, and Zaprionus indianus. 4. Exotic drosophilids breed in more plant species than Neotropical drosophilids and use exotic hosts more frequently, possibly because they are generalists that have survived the trial of introduction and establishment in the Neotropics. Native drosophilids are more variable in resource breadth and sometimes adopt exotic hosts. 5. Amongst the 49 drosophilids with enough records for analysis (> 4), 48 were categorised as generalists. One possible explanation for such overwhelming generalism is the high diversity of Neotropical habitat or hosts. A second, non‐exclusive explanation, suggested by recent studies and empirically supported by the absence of host specialisation found in this study, is that drosophilids could be selective of the dominant yeasts and bacteria in host tissue, and not of the hosts themselves.     

Use of an exotic host plant affects mate choice in an insect herbivore

The colonization of exotic plants by herbivorous insects has provided opportunities for investigating causes and consequences of the evolution of niche breadth. The butterfly Lycaeides melissa utilizes exotic alfalfa, Medicago sativa, which is a relatively poor larval resource, and previous studies have found that caterpillars that consume M. sativa develop into smaller and less fecund adults. Here we investigate the effect of smaller female body size on male mate preference, a previously unexplored consequence of novel host use. Smaller females, which developed on the exotic host, were less likely to be visited by males. This result was confirmed with a second set of choice tests involving females reared on a single plant species, thus ruling out host-specific confounding factors. We suggest that an effect on mate choice be considered part of the complex suite of factors determining persistence of herbivorous insects following colonization of new habitats or resources.     

Do Native Parasitic Plants Cause More Damage to Exotic Invasive Hosts Than Native Non-Invasive Hosts? An Implication for Biocontrol

Field studies have shown that native, parasitic plants grow vigorously on invasive plants and can cause more damage to invasive plants than native plants. However, no empirical test has been conducted and the mechanism is still unknown. We conducted a completely randomized greenhouse experiment using 3 congeneric pairs of exotic, invasive and native, non-invasive herbaceous plant species to quantify the damage caused by parasitic plants to hosts and its correlation with the hosts' growth rate and resource use efficiency. The biomass of the parasitic plants on exotic, invasive hosts was significantly higher than on congeneric native, non-invasive hosts. Parasites caused more damage to exotic, invasive hosts than to congeneric, native, non-invasive hosts. The damage caused by parasites to hosts was significantly positively correlated with the biomass of parasitic plants. The damage of parasites to hosts was significantly positively correlated with the relative growth rate and the resource use efficiency of its host plants. It may be the mechanism by which parasitic plants grow more vigorously on invasive hosts and cause more damage to exotic, invasive hosts than to native, non-invasive hosts. These results suggest a potential biological control effect of native, parasitic plants on invasive species by reducing the dominance of invasive species in the invaded community.     

Responses of a native beetle to novel exotic plant species with varying invasion history

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Temporal and geographic variation in parasitoid attack with no evidence for ant protection of the Melissa blue butterfly,Lycaeides melissa

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Trade-offs in larval performance on normal and novel hosts

The evolution of host specialization in phytophagous insects is generally thought to involve genetic trade-offs that prevent individuals from maximizing fitness simultaneously on two or more hosts. Several hypotheses, however, have suggested that trade-offs may not be evident in experiments comparing larval performance on normal and novel hosts. Tests on survivorship, growth rate, and pupal mass among families of the swallowtail butterfly Papilio oregonius on its normal host and on a novel host provide support for these hypotheses, although they do not discriminate among them. Families differed in their relative performance on the hosts, but there was no evidence of a negative genetic correlation between hosts for any of the measures of performance. In addition, there were no correlations among the different measures, corroborating an earlier result suggesting that these different components of performance in the P. machaon species group are under at least partially separate genetic control. These results and similar results published for other insects have now produced a body of studies indicating that genetic trade-offs in individual components of larval performance may not be a major factor preventing shifts onto novel host plants. Trade-offs leading to the evolution of host specialization are more likely to involve coordination among the various components of performance together with ecological factors that allow higher fitness on one host than on others.