Evolution Of Host Use Research Articles

Discrepancies between subgeneric classification and molecular phylogeny of Ceratitis (Diptera: Tephritidae), can the evolution of host use provide some clues?

Using molecular data from three protein encoding genes and 49 taxa (98 specimens from 20 African countries), we provide an extended phylogeny of Ceratitis and investigate the evolution of stenophagy across clades. Bayesian tree reconstructions support previously proposed monophyletic lineages ( Pardalaspis, Pterandrus section A, Pterandrus section B + Ceratitis sensu stricto) and reveal the occurrence of two new monophyletic groups including Ceratalaspis/ Hoplolophomyia ( viz. Cl(A), and Cl(B) + H). The reconstruction of ancestral character states shows that stenophagy evolved repeatedly and independently in five different clades ( Podocarpus, Solanum, Strychnos, Tabernaemontana and Vepris feeders). The evolution of feeding preferences is closely related to the phylogenetic patterns of Ceratalaspis/ Hoplolophomyia whose sections include either polyphagous species ( Cl(A)) or stenophagous taxa ( Cl(B) + H) that are further subdivided in Vepris and Solanum feeders. The evolution of stenophagy in the genus Ceratits appears as the result of a process leading to the exploitation of “unconventional” fruits ( viz. toxic and/or not fleshy) and involving either metabolic adaptation to toxic plant compounds and/or the capability of penetrating fruits with thick cuticles.

Can Preference for Oviposition Sites Initiate Reproductive Isolation in Callosobruchus maculatus?

Theory has identified a variety of evolutionary processes that may lead to speciation. Our study includes selection experiments using different host plants and test key predictions concerning models of speciation based on host plant choice, such as the evolution of host use (preference and performance) and assortative mating. This study shows that after only ten generations of selection on different resources/hosts in allopatry, strains of the seed beetle Callosobruchus maculatus develop new resource preferences and show resource-dependent assortative mating when given the possibility to choose mates and resources during secondary contact. The resulting reduced gene flow between the different strains remained for two generations after contact before being overrun by disassortative mating. We show that reduced gene flow can evolve in a population due to a link between host preference and assortative mating, although this result was not found in all lines. However, consistent with models of speciation, assortative mating alone is not sufficient to maintain reproductive isolation when individuals disperse freely between hosts. We conclude that the evolution of reproductive isolation in this system cannot proceed without selection against hybrids. Other possible factors facilitating the evolution of isolation would be longer periods of allopatry, the build up of local adaptation or reduced migration upon secondary contact.

Ecological interactions are evolutionarily conserved across the entire tree of life

Ecological interactions are crucial to understanding both the ecology and the evolution of organisms. Because the phenotypic traits regulating species interactions are largely a legacy of their ancestors, it is widely assumed that ecological interactions are phylogenetically conserved, with closely related species interacting with similar partners. However, the existing empirical evidence is inadequate to appropriately evaluate the hypothesis of phylogenetic conservatism in ecological interactions, because it is both ecologically and taxonomically biased. In fact, most studies on the evolution of ecological interactions have focused on specialized organisms, such as some parasites or insect herbivores, belonging to a limited subset of the overall tree of life. Here we study the evolution of host use in a large and diverse group of interactions comprising both specialist and generalist acellular, unicellular and multicellular organisms. We show that, as previously found for specialized interactions, generalized interactions can be evolutionarily conserved. Significant phylogenetic conservatism of interaction patterns was equally likely to occur in symbiotic and non-symbiotic interactions, as well as in mutualistic and antagonistic interactions. Host-use differentiation among species was higher in phylogenetically conserved clades, irrespective of their generalization degree and taxonomic position within the tree of life. Our findings strongly suggest a shared pattern in the organization of biological systems through evolutionary time, mediated by marked conservatism of ecological interactions among taxa.

Not All Oak Gall Wasps Gall Oaks: The Description of Dryocosmus rileypokei, A New, Apostate Species of Cynipini from California

Cynipini gall wasps (Hymenoptera: Cynipidae) are commonly known as oak gall wasps for their almost exclusive use of oak (Quercus spp.; Fagaceae) as their host plant. Previously, only three of the nearly 1,000 species of Cynipini have been recorded from hosts other than Quercus. These three are known from western chinquapin (Chrysolepis), chestnut (Castanea) and tan bark oak (Lithocarpus), all lineages of Fagaceae related to Quercus. Here we describe Dryocosmus rileypokei Morita & Buffington, new species, a second species of cynipine which attacks Chrysolepis. Unlike the previously known gall wasp D. castanopsidis, which produces a medium-sized spherical external gall near the base of the staminate (male) flowers of Chrysolepis sempervirens, D. rileypokei attacks the same host acting as a nut galler. Dryocosmus rileypokei creates a gall within the mesocarp wall of the nut and appears to draw nutrients away from the developing seed. Later instar larvae and teneral adults were found within these internal galls. It appears that the adult wasp eventually chews an exit hole from these galleries. The evolution of host use in the three, non-oak galling Dryocosmus species is discussed.

PHYLOGENETIC AND GEOGRAPHIC VARIATION IN HOST BREADTH AND COMPOSITION BY HERBIVOROUS AMPHIPODS IN THE FAMILY AMPITHOIDAE

Predicting the host range for herbivores has been a major aim of research into plant-herbivore interactions and an important model system for understanding the evolution of feeding specialization. Among many terrestrial insects, host range is strongly affected by herbivore phylogeny and long historical associations between particular herbivore and plant taxa. For small herbivores in marine environments, it is known that the evolution of host use is sculpted by several ecological factors (e.g., food quality, value as a refuge from predators, and abiotic forces), but the potential for phylogenetic constraints on host use remains largely unexplored. Here, we analyze reports of host use of herbivorous amphipods from the family Ampithoidae (102 amphipod species from 12 genera) to test the hypotheses that host breadth and composition vary among herbivore lineages, and to quantify the extent to which nonpolar secondary metabolites mediate these patterns. The family as a whole, and most individual species, are found on a wide variety of macroalgae and seagrasses. Despite this polyphagous host use, amphipod genera consistently differed in host range and composition. As an example, the genus Peramphithoe rarely use available macrophytes in the order Dictyotales (e.g., Dictyota) and as a consequence, display a more restricted host range than do other genera (e.g., Ampithoe, Cymadusa, or Exampithoe). The strong phylogenetic effect on host use was independent of the uneven distribution of host taxa among geographic regions. Algae that produced nonpolar secondary metabolites were colonized by higher numbers of amphipod species relative to chemically poor genera, consistent with the notion that secondary metabolites do not provide algae an escape from amphipod herbivory. In contrast to patterns described for some groups of phytophagous insects, marine amphipods that use chemically rich algae tended to have broader, not narrower, host ranges. This result suggests that an evolutionary advantage to metabolite tolerance in marine amphipods may be that it increases the availability of appropriate algal hosts (i.e., enlarges the resource base).

Phylogenetic relationships within the leaf-mining flies (Diptera: Agromyzidae) inferred from sequence data from multiple genes

The leaf-mining flies (Diptera: Agromyzidae) are a diverse group whose larvae feed internally in leaves, stems, flowers, seeds, and roots of a wide variety of plant hosts. The systematics of agromyzids has remained poorly known due to their small size and morphological homogeneity. We investigated the phylogenetic relationships among genera within the Agromyzidae using parsimony and Bayesian analyses of 2965 bp of DNA sequence data from the mitochondrial COI gene, the nuclear ribosomal 28S gene, and the single copy nuclear CAD gene. We included 86 species in 21 genera, including all but a few small genera, and spanning the diversity within the family. The results from parsimony and Bayesian analyses were largely similar, with major groupings of genera in common. Specifically, both analyses recovered a monophyletic Phytomyzinae and a monophyletic Agromyzinae. Within the subfamilies, genera found to be monophyletic given our sampling include Agromyza, Amauromyza, Calycomyza, Cerodontha, Liriomyza, Melanagromyza, Metopomyza, Nemorimyza, Phytobia, and Pseudonapomyza. Several genera were found to be polyphyletic or paraphyletic including Aulagromyza, Chromatomyia, Phytoliriomyza, Phytomyza, and Ophiomyia. We evaluate our findings and discuss host-use evolution in light of current agromyzid taxonomy and two recent hypotheses of relationships based on morphological data.

Pollen-host specificity and evolutionary patterns of host switching in a clade of specialist bees (Apoidea: Diadasia)

We examined the levels of pollen-host specificity in North American Diadasia (Hymenoptera: Apoidea), a clade of specialist bees. We analysed the scopal pollen loads of 409 individuals representing 25 of the 30 species of Diadasia that occur in North America. Each Diadasia species showed a preference for one of five plant families. However, the 25 species varied in their level of host specificity: the average percentage by volume of the preferred host in pollen loads ranged from > 99% to < 75%. In 17 of the 25 species, all or most individuals examined contained pure loads of one host taxon, while in eight species individuals were less specialized and carried mixtures of several unrelated host taxa. Mapping these host preferences onto a phylogenetic tree indicated that Malvaceae is the most likely ancestral host for the genus, and use of other hosts can be explained by a single switch to each of the other four host-plant families. Thus, most speciation events were not associated with a host switch; this pattern does not support host switching as a niche partitioning strategy to avoid competition. Diadasia species are more likely to use host-plant families that are used by other Diadasia and Emphorine bees; however, there was no evidence of residual adaptation to ancestral hosts. Diet breadth appears to be a labile trait: transitions from narrower to broader host use, as well as vice versa, were observed. The observed patterns of host-use evolution may be driven, in part, by host morphology and/or chemistry. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 487‐ 505.

Isolation and characterization of polymorphic microsatellite markers in Zabrotes subfasciatus Boheman (Coleoptera: Bruchidae)

AbstractBruchid beetles of the genus Zabrotes (Coleoptera: Bruchidae) are important worldwide pests of legume seeds. Samples of Zabrotes subfasciatus and Z. sylvestris were collected from seeds of two Phaseolus species throughout Mexico to assess the role of host plant variation and plant domestication on the evolution of host use in this species. For this purpose six polymorphic microsatellite loci were isolated and characterized for the bruchid Z. subfasciatus. Cross‐species amplification tests were performed on Z. sylvestris and revealed that three loci amplified successfully and were polymorphic in this closely related species.

A comparative study of interspecies mating of Phratora vulgatissima and P. vitellinae using behavioural tests and molecular markers

AbstractThe leaf beetle genus Phratora L. (Coleoptera: Chrysomelidae) has been used to study the ecology of host plant chemicals in herbivore preference, and the evolution of host use in chemical defence. Phratora vulgatissima and P. vitellinae are sympatric species distributed widely across Europe. Their trophic niches are largely separate due to strong differences in their host feeding preference, but they have occasionally been recorded together, feeding on Salix burjatica‘Germany’ and, only in early spring, on Populus trichocarpa (Torr &amp; A. Gray) ‘Trichobel’. Using behavioural tests and recently developed species‐specific microsatellite markers, the intra‐ and interspecific mating of both beetle species were investigated. The microsatellite markers provided evidence that interspecific mating occurred under field conditions. Interspecific mating also took place under laboratory conditions, but less frequently than mating within species. Females of both species laid fewer eggs, and fewer eggs per clutch, when isolated with an interspecific male than with a conspecific male. Female P. vulgatissima were polyandrous, as microsatellite markers showed that their larvae were the progeny of both P. vulgatissima males that had been isolated with a single female. While only 0.55% of eggs laid in interspecific pair combinations hatched, microsatellite markers provided evidence of hybridisation between beetle species; however, these larvae died within a week when reared in a Petri dish containing ‘Germany’ and P. trichocarpa leaves. It can therefore be inferred that reproductive isolation is complete. The results are discussed in relation to species integrity and the implications for diverse mixtures of short‐rotation coppice willow plantations.

Macroevolution of insect-plant associations: the relevance of host biogeography to host affiliation.

Identifying the factors that have promoted host shifts by phytophagous insects at a macroevolutionary scale is critical to understanding the associations between plants and insects. We used molecular phylogenies of the beetle genus Blepharida and its host genus Bursera to test whether these insects have been using hosts with widely overlapping ranges over evolutionary time. We also quantified the importance of host range coincidence relative to host chemistry and host phylogenetic relatedness. Overall, the evolution of host use of these insects has not been among hosts that are geographically similar. Host chemistry is the factor that best explains their macroevolutionary patterns of host use. Interestingly, one exceptional polyphagous species has shifted among geographically close chemically dissimilar plants.

Genetic basis for established and novel host plant use in a herbivorous ladybird beetle, Epilachna vigintioctomaculata

AbstractGenetic trade‐offs in host plant use are thought to promote the evolution of host specificity. Experiments on a range of herbivorous insects, however, have found negative genetic correlation in host plant use in only a limited number of species. To account for the general lack of negative genetic correlation, recent hypotheses advocate that different stages in evolution of host use must be distinguished: initial performance on a novel host in comparison with the established host, and performance on both hosts after the insect population has interacted with both hosts for a long time. The hypotheses suggest that genetic correlation may not necessarily be negative at the initial stage. The present study examines growth performance on both the established and a novel host in a herbivorous ladybird beetle, Epilachna vigintioctomaculata Motschulsky (Coccinellidae, Epilachninae). The results show that traits of growth performance across hosts were positively or neutrally correlated, but there was no evidence of a negative genetic correlation. In addition, significant genetic variance of growth performance on each host was detected, suggesting that E. vigintioctomaculata can potentially respond to selection for increased performance on both plant species. These results and similar results from experiments on other herbivores suggest that host expansion may not be constrained genetically, at least at the initial stage of host range evolution.

Comparative Detoxification of Plant ( Magnolia virginiana ) Allelochemicals by Generalist and Specialist Saturniid Silkmoths

The foliage of sweetbay magnolia (Magnolia virginana) contains at least two biologically active phenylpropanoid compounds (magnolol and a biphenyl ether) that are toxic to a number of generalist insect herbivores. These compounds have little effect on caterpillars of the sweetbay silkmoth, C. securifera, which is a specialist on sweetbay, but they are toxic to two closely related silkmoths, C. angulifera and C. promethea. To understand the influence of phytochemistry on the evolution of host use and feeding specialization in Callosamia, the detoxification capability of C. securifera was compared with that of C. angulifera and C. promethea. Degradation of magnolol and the biphenyl ether by midgut homogenate of the sweetbay specialist was NADPH-dependent and inhibited by piperonyl butoxide, suggesting the involvement of cytochrome P-450 detoxification enzymes. Both were degraded three times faster in the specialist compared to the unadapted herbivores. Higher rates of degradation could not be induced in the polyphagous C. promethea by a mixture of magnolol and the biphenyl ether or by the P-450 inducer pentamethylbenzene, nor did activity vary significantly when larvae were reared on different host plants. Use of sweetbay by Callosamia silkmoths appears to be dependent on their ability to degrade host toxins rapidly via midgut detoxification enzymes. Moreover, the intraspecific comparisons contradict the common prediction that higher levels of cytochrome P-450 activity are found in more polyphagous species; instead, P-450 activity is more closely associated with specific chemical attributes of the herbivores' host plants.

IS SPECIALIZATION A DEAD END? THE PHYLOGENY OF HOST USE IN DENDROCTONUS BARK BEETLES (SCOLYTIDAE).

Ecological explanations for the prevalence of resource specialists are abundant, whereas phylogenetic evidence on their origins is scarce. In this paper, we provide a molecular phylogenetic study of the 19 specialist or generalist species in the bark beetle genus Dendroctonus, which collectively attack species in four different genera in the conifer family Pinaceae. Given substantial variation in diet breadth, we asked two general questions concerning the evolution of resource use in this group. How conservative is the evolution of host use in these insects? Does specialization tend to be derived (i.e., a "dead end")? To answer these questions, we estimated the phylogeny of Dendroctonus using mitochondrial DNA sequences and mapped transitions in resource use on the resulting phylogeny estimate. The evolution of affiliations with Pinus and Picea hosts in Dendroctonus was conservative among beetle species (PTP test; P < 0.012), but there was no significant correspondence between the phylogeny of these beetles and the phylogeny among their Pinaceae hosts (among genera, P = 0.28; among Pinus species, P = 0.82). Degree of specialization, as measured in the proportion of hosts used, was bimodally distributed with "generalist" species utilizing < 60% of the congeneric hosts within their range and six specialist species utilizing < 40% of the available hosts. Among the generalists, we found a strong correlation between the number of hosts encountered and the number of hosts utilized (R = 0.97, P < 0.0001), whereas there was no significant correlation among the specialists (R = 0.27, P = 0.59). The evolution of specialization in Dendroctonus proved highly labile-specialists arose from generalists at least six separate times (without reversal) all in derived positions, and closer examination of some specialists revealed instances where they appear to have lost particular host species from their diet. However, evidence from the ecological literature also suggests that several Dendroctonus generalists may have increased their range of host genera within the Pinaceae.

Evolution of host use in fruit flies of the genus Blepharoneura (Diptera: Tephritidae): cryptic species on sexually dimorphic host plants

Abstract Blepharoneura , a neotropical genus of tephritid fruit flies, includes newly discovered morphologically cryptic species that are highly host specific. Several species of fly, each specific to a different host tissue, can infest a single species of sexually dimorphic host. Using allozyme electrophoresis, we analysed morphologically similar populations of flies reared from various tissues of five species of host plant representing three subtribes of the Cucurbitaceae. In Venezuela, we found three sympatric species of Blepharoneura feeding on Gurania spinulosa Cogn. (one species on male flowers, another on female flowers, and a third on seeds). In Costa Rica, we found four non-interbreeding populations of Blepharoneura feeding on G. costaricensis Cogn.: two sympatric «lowland» populations (one feeding on male flowers and another on female flowers) and two sympatric «highland» populations (also specific to flowers of different sexes). Phylogenetic analyses revealed a species-rich clade of specialists associated with flowers of the Guraniinae, and a species-poor clade of generalists associated with seeds of the Guraniinae.Specialization on male vs. female flowers appears to have occurred repeatedly. Patterns of larval dispersal may underlie differences in diversification of seed vs. flower clades: larvae that feed on seeds are dispersed by bats; larvae feeding on flowers are not dispersed.

Evolution of host use in fruit flies of the genusBlepharoneura(Diptera: Tephritidae): cryptic species on sexually dimorphic host plants

Blepharoneura, a neotropical genus of tephritid fruit flies, includes newly discovered morphologically cryptic species that are highly host specific. Several species of fly, each specific to a different host tissue, can infest a single species of sexually dimorphic host. Using allozyme electrophoresis, we analysed morphologically similar populations of flies reared from various tissues of five species of host plant representing three subtribes of the Cucurbitaceae. In Venezuela, we found three sympatric species ofBlepharoneurafeeding onGurania spinulosaCogn. (one species on male flowers, another on female flowers, and a third on seeds). In Costa Rica, we found four non-interbreeding populations ofBlepharoneurafeeding onG. costaricensisCogn.: two sympatric «lowland» populations (one feeding on male flowers and another on female flowers) and two sympatric «highland» populations (also specific to flowers of different sexes). Phylogenetic analyses revealed a species-rich clade of specialists associated with flowers of the Guraniinae, and a species-poor clade of generalists associated with seeds of the Guraniinae.Specialization on male vs. female flowers appears to have occurred repeatedly. Patterns of larval dispersal may underlie differences in diversification of seed vs. flower clades: larvae that feed on seeds are dispersed by bats; larvae feeding on flowers are not dispersed.