Evolutionary dynamics of host-plant specialization: a case study of the tribe Nymphalini

Two general patterns that have emerged from the intense studies on insect-host plant associations are a predominance of specialists over generalists and a taxonomic conservatism in host-plant use. In most insect-host plant systems, explanations for these patterns must be based on biases in the processes of host colonizations, host shifts, and specialization, rather than cospeciation. In the present paper, we investigate changes in host range in the nymphalid butterfly tribe Nymphalini, using parsimony optimizations of host-plant data on the butterfly phylogeny. In addition, we performed larval establishment tests to search for larval capacity to feed and survive on plants that have been lost from the female egg-laying repertoire. Optimizations suggested an ancestral association with Urticaceae, and most of the tested species showed a capacity to feed on Urtica dioica regardless of actual host-plant use. In addition, there was a bias among the successful establishments on nonhosts toward plants that are used as hosts by other species in the Nymphalini. An increased likelihood of colonizing ancestral or related plants could also provide an alternative explanation for the observed pattern that some plant families appear to have been colonized independently several times in the tribe. We also show that there is no directionality in host range evolution toward increased specialization, that is, specialization is not a dead end. Instead, changes in host range show a very dynamic pattern.     

Diversity and evolution of ectomycorrhizal host associations in the Sclerodermatineae (Boletales, Basidiomycota)

This study uses phylogenetic analysis of the Sclerodermatineae to reconstruct the evolution of ectomycorrhizal host associations in the group using divergence dating, ancestral range and ancestral state reconstructions. Supermatrix and supertree analysis were used to create the most inclusive phylogeny for the Sclerodermatineae. Divergence dates were estimated in BEAST. Lagrange was used to reconstruct ancestral ranges. BayesTraits was used to reconstruct ectomycorrhizal host associations using extant host associations with data derived from literature sources. The supermatrix data set was combined with internal transcribed spacer (ITS) data sets for Astraeus, Calostoma, and Pisolithus to produce a 168 operational taxonomic unit (OTU) supertree. The ensuing analysis estimated that basal Sclerodermatineae originated in the late Cretaceous while major genera diversified near the mid Cenozoic. Asia and North America are the most probable ancestral areas for all Sclerodermatineae, and angiosperms, primarily rosids, are the most probable ancestral hosts. Evolution in the Sclerodermatineae follows the biogeographic history of disjunct plant communities associated with early Cenozoic mesophytic forests and a boreotropical history. Broad geographic distributions are observed in the most promiscuous Sclerodermatineae (those with broad host ranges), while those with relatively limited distribution have fewer documented ectomycorrhizal associations. This suggests that ectomycorrhizal generalists have greater dispersal capabilities than specialists.     

Host shifts in economically significant fruit flies (Diptera: Tephritidae) with high degree of polyphagy

Insects tend to feed on related hosts. Coevolution tends to be dominated by interactions resulting from plant chemistry in defense strategies, and evolution of secondary metabolisms being in response to insect herbivory remains a classic explanation of coevolution. The present study examines whether evolutionary constraints existing in host associations of economically important fruit flies in the species‐rich tribe Dacini (Diptera: Tephritidae) and to what extent these species have evolved specialized dietary patterns. We found a strong effect of host phylogeny on associations on the 37 fruit flies tested, although the fruit fly species feeding on ripe commercially grown fruits that lost the toxic compounds after long‐term domestication are mostly polyphagous. We assessed the phylogenetic signal of host breadth across the fruit fly species, showing that the results were substantially different depending on partition levels. Further, we mapped main host family associations onto the fruit fly phylogeny and Cucurbitaceae has been inferred as the most likely ancestral host family for Dacini based on ancestral state reconstruction.     

The phylogenetics and biochemistry of host-plant specialization in Melitaeine butterflies (Lepidoptera: Nymphalidae)

Butterflies in the tribe Melitaeini (Lepidoptera: Nymphalidae) are known to utilize host plants belonging to 16 families, although most host-plant records are from four families. Of the 16 host-plant families, 12 produce secondary plant metabolites called iridoids. Earlier studies have shown that larvae of several melitaeine species use iridoids as feeding stimulants and sequester these compounds for larval defense. I investigate the evolutionary history of host-plant use in the tribe Melitaeini by testing a recent phylogenetic hypothesis of 65 species representing the four major species groups of the tribe. By simple character optimization of host-plant families and presence/absence of iridoids in the host plants, I find that plant chemistry is a more conservative trait than plant taxonomy. The ancestral host plant(s) of the entire tribe most likely contained iridoids and were likely to be in the plant family Plantaginaceae. A major host shift from plants containing iridoids to plants not containing iridoids has happened three times independently. The results show that the evolution of host-plant use in melitaeines has been (and still is) a dynamic process when considering plant taxonomy, but is relatively stable when considering host-plant chemistry.     

Experimentally induced host‐shift changes life‐history strategy in a seed beetle

Expansion of the host range in phytophagous insects depends on their ability to form an association with a novel plant through changes in host‐related traits. Phenotypic plasticity has important effects on initial survival of individuals faced with a new plant, as well as on the courses of evolutionary change during long‐term adaptation to novel conditions. Using experimental populations of the seed beetle that evolved on ancestral (common bean) or novel (chickpea) host and applying reciprocal transplant at both larval and adult stage on the alternative host plant, we studied the relationship between the initial (plastic) phases of host‐shift and the subsequent stages of evolutionary divergence in life‐history strategies between populations exposed to the host‐shift process. After 48 generations, populations became well adapted to chickpea by evolving the life‐history strategy with prolonged larval development, increased body mass, earlier reproduction, shorter lifespan and decreased plasticity of all traits compared with ancestral conditions. In chickpea‐adapted beetles, negative fitness consequences of low plasticity of pre‐adult development (revealed as severe decrease in egg‐to‐adult viability on beans) exhibited mismatch with positive effects of low plasticity (i.e. low host sensitivity) in oviposition and fecundity. In contrast, beetles adapted to the ancestral host showed high plasticity of developmental process, which enabled high larval survival on chickpea, whereas elevated plasticity in adult behaviour (i.e. high host sensitivity) resulted in delayed reproduction and decreased fecundity on chickpea. The analysis of population growth parameters revealed significant fluctuation during successive phases of the host‐shift process in A. obtectus.     

BUTTERFLIES AND PLANTS: A PHYLOGENETIC STUDY

A database on host plant records from 437 ingroup taxa has been used to test a number of hypotheses on the interaction between butterflies and their host plants using phylogenetic methods (simple character optimization, concentrated changes test, and independent contrasts test). The butterfly phylogeny was assembled from various sources and host plant clades were identified according to Chase et al.'s rbcL-based phylogeny. The ancestral host plant appears to be associated within a highly derived rosid clade, including the family Fabaceae. As fossil data suggest that this clade is older than the butterflies, they must have colonized already diversified plants. Previous studies also suggest that the patterns of association in most insect-plant interactions are more shaped by host shifts, through colonization and specialization, than by cospeciation. Consequently, we have focused explicitly on the mechanisms behind host shifts. Our results confirm, in the light of new phylogenetic evidence, the pattern reported by Ehrlich and Raven that related butterflies feed on related plants. We show that host shifts have generally been more common between closely related plants than between more distantly related plants. This finding, together with the possibility of a higher tendency of recolonizing ancestral hosts, helps to explain the apparent large-scale conservation in the patterns of association between insects and their host plants, patterns which at the same time are more flexible on a more detailed level. Plant growth form was an even more conservative aspect of the interaction between butterflies and their host plants than plant phylogeny. However, this is largely explained by a higher probability of colonizations and host shifts while feeding on trees than on other growth forms.     

The evolution of generalization? Parasitoid flies and the perils of inferring host range evolution from phylogenies

It is widely assumed that high resource specificity predisposes lineages toward greater likelihood of extinction and lower likelihood of diversification than more generalized lineages. This suggests that host range evolution in parasitic organisms should proceed from generalist to specialist, and specialist lineages should be found at the 'tips' of phylogenies. To test these hypotheses, parsimony and maximum likelihood methods were used to reconstruct the evolution of host range on a phylogeny of parasitoid flies in the family Tachinidae. In contrast to predictions, most reconstructions indicated that generalists were repeatedly derived from specialist lineages and tended to occupy terminal branches of the phylogeny. These results are critically examined with respect to hypotheses concerning the evolution of specialization, the inherent difficulties in inferring host ranges, our knowledge of tachinid-host associations, and the methodological problems associated with ancestral character state reconstruction. Both parsimony and likelihood reconstructions are shown to provide misleading results and it is argued that independent evidence, in addition to phylogenetic trees, is needed to inform models of the evolution of host range and the evolutionary consequences of specialization.     

A new species Megabruchidius sophorae (Coleoptera, Bruchidae), feeding on seeds of Styphnolobium (Fabaceae) new to Bruchidae

A new species Megabruchidiussophorae (Insecta, Coleoptera) is described from Japan (Honshu). The larval host of this bruchid is the seeds of the tree legume 'enju', or chinese scholar tree, Styphnolobium japonicum (a senior synonym of Sophora japonica), which is a new host genus to Bruchidae. Styphnolobium is positioned basally in molecular phylogeny of the leguminous subfamily Papilionoideae. Other members of Megabruchidius are known to feed on Gleditsia, the tree legumes that belong to the most ancestral subfamily Caesalpinioideae. Therefore, Megabruchidius utilizes ancestral groups of legumes as its host plants. Megabruchidius has been inferred to be ancestral, based on its behavior. The character state of the host for this third Megabruchidius species supports that the genus is ancestral, at least in the subfamily Bruchinae. We also reviewed the genera closely related to Megabruchidius, i.e., Bruchidius and Sulcobruchus in Bruchidini, and wrote a key to the species in the genus Megabruchidius.     

Macroevolutionary patterns in the Aphidini aphids (Hemiptera: Aphididae): diversification, host association, and biogeographic origins

Background

Due to its biogeographic origins and rapid diversification, understanding the tribe Aphidini is key to understanding aphid evolution. Major questions about aphid evolution include origins of host alternation as well as age and patterns of diversification in relation to host plants. To address these questions, we reconstructed the phylogeny of the Aphidini which contains Aphis, the most diverse genus in the family. We used a combined dataset of one nuclear and four mitochondrial DNA regions. A molecular dating approach, calibrated with fossil records, was used to estimate divergence times of these taxa.

Principal Findings

Most generic divergences in Aphidini occurred in the Middle Tertiary, and species-level divergences occurred between the Middle and Late Tertiary. The ancestral state of host use for Aphidini was equivocal with respect to three states: monoecy on trees, heteroecy, and monoecy on grasses. The ancestral state of Rhopalosiphina likely included both heteroecy and monoecy, whereas that of Aphidina was most likely monoecy. The divergence times of aphid lineages at the generic or subgeneric levels are close to those of their primary hosts. The species-level divergences in aphids are consistent with the diversification of the secondary hosts, as a few examples suggest. The biogeographic origin of Aphidini as a whole was equivocal, but the major lineages within Aphidina likely separated into Nearctic, Western Palearctic, and Eastern Palearctic regions.

Conclusions

Most generic divergences in Aphidini occurred in the Middle Tertiary when primary hosts, mainly in the Rosaceae, were diverging, whereas species-level divergences were contemporaneous with diversification of the secondary hosts such as Poaceae in the Middle to Late Tertiary. Our results suggest that evolution of host alternation within Aphidini may have occurred during the Middle Tertiary (Oligocene) when the secondary hosts emerged.     

Ancient association with Fagaceae in the aphid tribe Greenideini (Hemiptera: Aphididae: Greenideinae)

Aphids are intimately associated with their host plants. Evolutionary lability of host association is common within heteroecious aphid lineages, whereas our knowledge of host‐use evolution in non‐host‐alternating aphids is limited. In the present study, we construct the first detailed molecular phylogeny of the monoecious aphid tribe Greenideini based on three mitochondrial genes (COI, COII and Cytb) and one nuclear gene (EF‐1α), and investigated its history of host association. Maximum likelihood and Bayesian phylogenies strongly support the monophyly of Greenideini and most constituent genera. Divergence time estimates and character reconstructions suggest that Greenideini may have originated during the Late Cretaceous to early Paleogene, which accompanies the origin of its ancestral host, members of the family Fagaceae. Colonisation of novel host plants has occurred multiple times during the evolutionary history of Greenideini, thereby leading to current patterns of host association. We suggest that directly shifting to novel hosts, together with expanding host range onto pre‐existing, unused plants, has probably promoted diversification in this tribe.     

Phylogeny and life-history evolution in Carabus (subtribe Carabina: Coleoptera, Carabidae) based on sequences of two nuclear genes

We reconstructed the phylogeny of the holarctic carabid subtribe Carabina (= Carabus s . l. ; Coleoptera, family Carabidae, supertribe Carabitae, tribe Carabini) using sequence data from two nuclear DNA loci: wingless ( Wg ) and phosphoenolpyruvate carboxykinase ( PepCK ). The analysis of the combined data resulted in a well-resolved tree, although Wg and PepCK had partially conflicting phylogenetic signals. The tree topology did not support the reciprocal monophyly of some previously proposed major divisions, whereas all but one of eight other previously proposed subdivisions were recovered as monophyletic clades; the exception was paraphyletic. When two key life-history traits, seasonal life cycle and larval diet, were mapped onto the combined tree, spring breeding (with no larval overwintering) and insectivorous were ancestral traits. Malacophagy has apparently evolved at least twice: once in a small basal group (ten species) and again in a large derived group ( c . 340 species) that accounts for 42% of the species of Carabina. A third type of larva, earthworm feeders, appeared in two related, derived clades and represented moderate species diversity (12%). From the ancestral spring-breeder type with no larval overwintering, autumn-breeder types with larval overwintering apparently evolved repeatedly in insectivorous and malacophagous species, but not in earthworm feeders. Therefore, the extant diversity of the Carabina depends on the high rate of differentiation in the most derived group of malacophagous species.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 135–149.     

Fritillary phylogeny, classification, and larval host plants: reconstructed mainly on the basis of male and female genitalic morphology (Lepidoptera: Nymphalidae: Argynnini)

Species-level phylogeny of the Butterfly tribe Argynnini is established based on 141 characters derived from wing and genitalia morphology of both sexes. The Argynnini can be divided into three subtribes; Yrameina comprising Yramea and Boloria s . l ., Argynnina comprising Prokuekenthaliella , Issoria , Brenthis , and all the 'large fritillary' species joined in the genus Argynnis s . l . and a new subtribe Euptoietina comprising only the genus Euptoieta . The classical genus Issoria s . l . is polyphyletic regarding Yramea and possibly paraphyletic regarding the two Afrotropic species baumanni and hanningtoni ; these two species are tentatively transferred to the old genus/subgenus Prokuekenthaliella . Surprisingly, one Afrotropic species, Issoria smaragdifera is closely related to the East Palaearctic Issoria species. A revised classification of Argynnini is proposed based on the obtained phylogeny. Studies of larval host plants based on the obtained phylogeny suggest that the ancestral Argynnini used Passiflora and Violaceae, but already the ancestor of Yrameina + Argynnina was probably specialized on Violaceae. Whereas the Boloria species have turned to other food plants such as Dryas , Vaccinium and Salix on several occasions, only Brenthis among the Argynnina use other host plants than Viola (mainly Rosaceae). The habit of laying eggs away from the food plant has probably evolved twice within Argynnina.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 627–673.     

Phylogenetic and ecological relationships of the Hawaiian Drosophila inferred by mitochondrial DNA analysis

The Hawaiian Drosophilidae are comprised of an estimated 1000 species, all arising from a single common ancestor in the last 25 million years. This group, because of its species diversity, marked sexual dimorphism and complex mating behavior, host plant specificity, and the well-known chronology of the Hawaiian Archipelago, is an excellent model system for evolutionary studies. Here we present a phylogeny of this group based on ~2.6 kb of mitochondrial DNA sequence. Our taxon sampling is the most extensive to date, with nearly 200 species representing all species groups and most subgroups from the larger clades. Our results suggest that the picture wing and modified mouthpart species, long believed to be derived within this radiation, may actually occupy a basal position in the phylogeny. The haleakale species group, in contrast, is strongly supported as sister to the AMC clade. We use the phylogenetic results to examine the evolution of two important ecological characters, the host family and type of substrate used for oviposition and larval development. Although both host and substrate transitions are common in the group, oviposition substrate is more conserved among species groups than host plant family. While the ancestral host plant family is equivocally reconstructed, our results suggest that the ancestor of this group may have used rotting bark as a primary oviposition substrate.     

Relationship between adult and larval host plant selection and larval performance in the generalist moth, Trichoplusia ni

Adult oviposition preferences are expected to correlate with host plant suitability for the development of their offspring. For most lepidopteran species, this is particularly important as the hatching neonate larvae of many species are relatively immobile. Thus, the site of oviposition chosen by a female adult can greatly influence the probability of survival for her offspring. In the present study, we investigated the oviposition preference of adult Trichoplusia ni moths for six plant species to determine whether they could accurately rank the suitability of the plants for larval development. We also compared oviposition preferences to neonate larval acceptance and preference to determine whether the adult host range matched that of larval diet breath. Our results indicate that in two-choice and no-choice tests adult T. ni were able to rank the plants accurately, with the exception of anise hyssop. However, when given a choice of all six plants together, they laid more eggs on a plant that was not suitable for larval survival. Larvae accepted and fed on all plants in no-choice tests, and accurately ranked them according to larval performance. We conclude that neonate larvae are better able than adults to rank plants according to larval performance, and that larval diet breadth is wider than the range of plants accepted by adults. We also provide a discussion of the reduced accuracy of adult oviposition preference with increased plant choices.     

Biology of Asphondyliini (Diptera: Cecidomyiidae)

The family Cecidomyiidae (Diptera) including about 6100 described species displays diverse feeding habits. The tribe Asphondyliini is a well‐circumscribed monophyletic group of Cecidomyiidae and all species are known as gall inducers. Species belonging to this tribe exhibit fascinating ecological traits such as host alternation, polyphagy, extended diapause, induction of dimorphic galls and association with fungal symbionts. For these reasons, biogeographical and phylogenetic studies of Asphondyliini are of interest in elucidating the evolution of these traits, and particularly the processes of host‐range expansion, host‐plant shift and shifts in gall‐bearing organs. In order to facilitate further evolutionary studies of Asphondyliini, I review studies of systematics, biogeography, phylogeny, speciation, cytology, behavior, ecology, physiology, biological interaction and economic importance in this tribe.     

Higher-level phylogeny of the Ithomiinae (Lepidoptera: Nymphalidae): classification, patterns of larval hostplant colonization and diversification

We present a higher‐level phylogenetic hypothesis for the diverse neotropical butterfly subfamily Ithomiinae, inferred from one of the largest non‐molecular Lepidoptera data sets to date, including 106 species (105 ingroup) and 353 characters (306 informative) from adult and immature stage morphology and ecology. Initial analyses resulted in 1716 most parsimonious trees, which were reduced to a single tree after successive approximations character weighting. The inferred phylogeny was broadly consistent with other past and current work. Although some deeper relationships are uncertain, tribal‐level clades were generally strongly supported, with two changes required to existing classification. The tribe Melinaeini is polyphyletic and Athesis + Patricia require a new tribe. Methona should be removed from Mechanitini into the restored tribe Methonini. Dircennini was paraphyletic in analyses of all data but monophyletic based on adult morphology alone, and its status remains to be confirmed. Hypothyris, Episcada, Godyris, Hypoleria and Greta are paraphyletic. A simulation analysis showed that relatively basal branches tended to have higher partitioned Bremer support for immature stage characters. Larval hostplant records were optimized on to a reduced, generic‐level phylogeny and indicate that ithomiines moved from Apocynaceae to Solanaceae twice, or that Tithoreini re‐colonized Apocynaceae after a basal shift to Solanaceae. Ithomiine clades have specialized on particular plant clades suggesting repeated colonization of novel hostplant niches consistent with adaptive radiation. The shift to Solanum, comprising 70% of neotropical Solanaceae, occurs at the base of a clade containing 89% of all ithomiines, and is interpreted as the major event in the evolution of ithomiine larval hostplant relationships. © The Willi Hennig Society 2006.     

Phylogeny of the parasitic wasp subfamily Euphorinae (Braconidae) and evolution of its host preferences

The braconid subfamily Euphorinae is a large, cosmopolitan group of endoparasitoid wasps. The majority of species attack adult hosts, a strategy that is rare among parasitic wasps, but there are also many species that attack nymphs and larval stages. Euphorine hosts may belong to a variety of insect orders (Coleoptera, Hemiptera, Hymenoptera, Neuroptera, Psocoptera, Orthoptera and Lepidoptera) although most euphorine tribes are confined to Coleoptera. Here we investigate the phylogenetic relationships of the Euphorinae based on molecular data (3 kb of nucleotide data from four markers: 18S, 28S, CAD and COI) and propose a higher‐level classification based upon the resulting phylogeny. We also infer the evolution of host associations and discuss the diversification of the Euphorinae. Results from both Bayesian inference and maximum‐likelihood analysis show that the subfamily, as previously circumscribed, is paraphyletic. We propose that the subfamily be expanded to include the tribes Meteorini and Planitorini (Mannokeraia + Planitorus), so that it corresponds to a clade that is strongly supported as monophyletic in our analyses. Based on our results, a revised higher classification of the Euphorinae is proposed, in which 52 extant genera and 14 tribes are recognized. We reinstate the genus Microctonus belonging to the tribe Perilitini, and synonymize Ussuraridelus with Holdawayella, Sinuatophorus with Eucosmophorus. Furthermore, we propose the following tribal rearrangements: Spathicopis and Stenothremma are transferred to Perilitini; Tuberidelus, Eucosmophorus and Plynops to Cosmophorini; Ecclitura to Dinocampini; Chrysopophthorus, Holdawayella and Wesmaelia to Helorimorphini; Proclithroporus and Heia to Townesilitini. The monotypic tribe Cryptoxilonini is synonymized with Cosmophorini. The genera Pygostolus and Litostolus are placed in a separate tribe, Pygostolini, previously recognized as a subtribe among the Centistini. Parsimony‐based ancestral state reconstructions suggest that the ancestor of Euphorinae was a parasitoid of lepidopteran larvae, and that a host shift to larval Coleoptera occurred only in one clade of the Meteorini, some members of which secondarily shifted back to larval lepidopteran hosts. In the remainder of the subfamily, there was an initial shift from larval to adult coleopterans, followed by subsequent shifts to adults or larvae of Hemiptera, Hymenoptera, Neuroptera, Orthoptera and Psocoptera.     

THE EVOLUTION OF AGRICULTURE IN BEETLES (CURCULIONIDAE: SCOLYTINAE AND PLATYPODINAE)

Abstract Beetles in the weevil subfamilies Scolytinae and Platypodinae are unusual in that they burrow as adults inside trees for feeding and oviposition. Some of these beetles are known as ambrosia beetles for their obligate mutualisms with asexual fungi—known as ambrosia fungi—that are derived from plant pathogens in the ascomycete group known as the ophiostomatoid fungi. Other beetles in these subfamilies are known as bark beetles and are associated with free‐living, pathogenic ophiostomatoid fungi that facilitate beetle attack of phloem of trees with resin defenses. Using DNA sequences from six genes, including both copies of the nuclear gene encoding enolase, we performed a molecular phylogenetic study of bark and ambrosia beetles across these two subfamilies to establish the rate and direction of changes in life histories and their consequences for diversification. The ambrosia beetle habits have evolved repeatedly and are unreversed. The subfamily Platypodinae is derived from within the Scolytinae, near the tribe Scolytini. Comparison of the molecular branch lengths of ambrosia beetles and ambrosia fungi reveals a strong correlation, which a fungal molecular clock suggests spans 60 to 21 million years. Bark beetles have shifted from ancestral association with conifers to angiosperms and back again several times. Each shift to angiosperms is associated with elevated diversity, whereas the reverse shifts to conifers are associated with lowered diversity. The unusual habit of adult burrowing likely facilitated the diversification of these beetle‐fungus associations, enabling them to use the biomass‐rich resource that trees represent and set the stage for at least one origin of eusociality.     

Host tree resistance against the polyphagous wood-boring beetle Anoplophora glabripennis

Anoplophora glabripennis (Motschulsky) (Coleoptera: Cerambycidae: Lamiini) is an invasive wood‐boring beetle with an unusually broad host range and a proven ability to increase its host range as it colonizes new areas and encounters new tree species. The beetle is native to eastern Asia and has become an invasive pest in North America and Europe, stimulating interest in delineating host and non‐host tree species more clearly. When offered a choice among four species of living trees in a greenhouse, adult A. glabripennis fed more on golden‐rain tree (Koelreuteria paniculata Laxmann) and river birch (Betula nigra L.) than on London planetree (Platanus × acerifolia (Aiton) Willdenow) or callery pear (Pyrus calleryana Decaisne). Oviposition rate was highest in golden‐rain tree, but larval mortality was also high and larval growth was slowest in this tree species. Oviposition rate was lowest in callery pear, and larvae failed to survive in this tree species, whether they eclosed from eggs laid in the trees or were manually inserted into the trees. Adult beetles feeding on callery pear had a reduced longevity and females feeding only on callery pear failed to develop any eggs. The resistance of golden‐rain tree against the larvae appears to operate primarily through the physical mechanism of abundant sap flow. The resistance of callery pear against both larvae and adults appears to operate through the chemical composition of the tree, which may include compounds that are toxic or which otherwise interfere with normal growth and development of the beetle. Unlike river birch or London planetree, both golden‐rain tree and callery pear are present in the native range of A. glabripennis and may therefore have developed resistance to the beetle by virtue of exposure to attack during their evolutionary history.     

Multiple origins of anural development in ascidians inferred from rDNA sequences

Ascidians exhibit two different modes of development. A tadpole larva is formed during urodele development, whereas the larval phase is modified or absent during anural development. Anural development is restricted to a small number of species in one or possibly two ascidian families and is probably derived from ancestors with urodele development. Anural and urodele ascidians constitute a model system in which to study the evolution of development, but the phylogeny of anural development has not been resolved. Classification based on larval characters suggests that anural species are monophyletic, whereas classification according to adult morphology suggests they are polyphyletic. In the present study, we have inferred the origin of anural development using rDNA sequences. The central region of 18S rDNA and the hypervariable D2 loop of 28S rDNA were amplified from the genomic DNA of anural and urodele ascidian species by the polymerase chain reaction and sequenced. Phylogenetic trees inferred from 18S rDNA sequences of 21 species placed anural developers into two discrete groups corresponding to the Styelidae and Molgulidae, suggesting that anural development evolved independently in these families. Furthermore, the 18S rDNA trees inferred at least four independent origins of anural development in the family Molgulidae. Phylogenetic trees inferred from the D2 loop sequences of 13 molgulid species confirmed the 18S rDNA phylogeny. Anural development appears to have evolved rapidly because some anural species are placed as closely related sister groups to urodele species. The phylogeny inferred from rDNA sequences is consistent with molgulid systematics according to adult morphology and supports the polyphyletic origin of anural development in ascidians. Correspondence to: W.R. Jeffery