Parallel diversification of Australian gall-thrips on Acacia

The diversification of gall-inducing Australian Kladothrips (Insecta: Thysanoptera) on Acacia has produced a pair of sister-clades, each of which includes a suite of lineages that utilize virtually the same set of 15 closely related host plant species. This pattern of parallel insect-host plant radiation may be driven by cospeciation, host-shifting to the same set of host plants, or some combination of these processes. We used molecular-phylogenetic data on the two gall-thrips clades to analyze the degree of concordance between their phylogenies, which is indicative of parallel divergence. Analyses of phylogenetic concordance indicate statistically-significant similarity between the two clades. Their topologies also fit with a hypothesis of some degree of host-plant tracking. Based on phylogenetic and taxonomic information regarding the phylogeny of the Acacia host plants in each clade, one or more species has apparently shifted to more-divergent Acacia host-plant species, and in each case these shifts have resulted in notable divergence in aspects of the phenotype including morphology, life history and behaviour. Our analyses indicate that gall-thrips on Australian Acacia have undergone parallel diversification as a result of some combination of cospeciation, highly restricted host-plant shifting, or both processes, but that the evolution of novel phenotypic diversity in this group is a function of relatively few shifts to divergent host plants. This combination of ecologically restricted and divergent radiation may represent a microcosm for the macroevolution of host plant relationships and phenotypic diversity among other phytophagous insects.     

Phylogenetic analyses of DNA and allozyme data suggest that Gonioctena leaf beetles (Coleoptera; Chrysomelidae) experienced convergent evolution in their history of host-plant family shifts

A phylogenetic analysis of the genus Gonioctena (Coleoptera, Chrysomelidae) based on allozyme data (17 loci) and mitochondrial DNA sequence data (three gene fragments, 1,391 sites) was performed to study the evolutionary history of host-plant shifts among these leaf beetles. This chrysomelid genus is characteristically associated with a high number of different plant families. The diverse molecular data gathered in this study are to a large extent congruent, and the analyses provide a well-supported phylogenetic hypothesis to address questions about the evolution of host-plant shifts in the genus Gonioctena. The most-parsimonious reconstruction of the ancestral host-plant associations, based on the estimated phylogeny, suggests that the Fabaceae was the ancestral host-plant family of the genus. Although most of the host-plant shifts (between different host species) in Gonioctena have occurred within the same plant family or within the same plant genus, at least eight shifts have occurred between hosts belonging to distantly related and chemically dissimilar plant families. In these cases, host shifts may have been simply directed toward plant species available in the environment. Yet, given that two Gonioctena lineages have independently colonized the same three new plant families (Salicaceae, Betulaceae, Rosaceae), including four of the same new genera (Salix, Alnus, Prunus, Sorbus), some constraints are likely to have limited the different possibilities of interfamilial host-plant shifts.     

Both host-plant phylogeny and chemistry have shaped the African seed-beetle radiation

For the last 40 years, many authors have attempted to characterize the main patterns of plant-insect evolutionary interactions and understand their causes. In the present work on African seed-beetles (Coleoptera: Bruchidae), we have performed a 10-year field work to sample seeds of more than 300 species of potential host-plants (from the family Fabaceae), to obtain bruchids by rearing. This seed sampling in the field was followed by the monitoring of adult emergences which gave us the opportunity to identify host-plant use accurately. Then, by using molecular phylogenetics (on a combined data set of four genes), we have investigated the relationships between host-plant preferences and insect phylogeny. Our objectives were to investigate the level of taxonomic conservatism in host-plant fidelity and host-plant chemistry. Our results indicate that phylogenetically related insects are associated with phylogenetically related host-plants but the phylogeny of the latter cannot alone explain the observed patterns. Major host shifts from Papilionoideae to Mimosoideae subfamilies have happened twice independently suggesting that feeding specialization on a given host-plant group is not always a dead end in seed-beetles. If host-plant taxonomy and chemistry in legumes generally provide consistent data, it appears that the nature of the seed secondary compounds may be the major factor driving the diversification of a large clade specializing on the subfamily Mimosoideae in which host-plant taxonomy is not consistent with chemical similarity.     

Parallels in the evolution of the two largest New and Old World seed-beetle genera (Coleoptera, Bruchidae)

This study provides the first phylogenetic analysis of a large sample of the two largest genera of seed-beetles, Acanthoscelides Schilsky and Bruchidius Schilsky, which mostly feed on legumes (Fabaceae). The goal of this study was to investigate evolutionary patterns in relation to biogeography and host-plant associations. We used three mitochondrial molecular markers and parsimony and Bayesian inference methods to reconstruct the phylogeny of 76 species. In addition, we critically reviewed host-plant records in the literature for these two bruchid genera. Our results demonstrated the existence of two major clades, one New World and one largely Old World, which generally correspond to the two genera. Yet, current classification of several species is erroneous, so that both genera as currently defined are paraphyletic. We highlighted a strong trend toward specialization (with high taxonomic conservatism in host-plant use) exhibited by the two studied genera. However, we showed the existence of several host shifts during the evolution of this group of bruchids. Our phylogenetic hypotheses and our evaluation of host-plant associations both suggest that the two genera have undergone parallel evolution, as they have independently colonized similar host plants in their respective areas of distribution. Our estimation of divergence times indicated a more ancient origin for bruchids than that suggested by the fossil records. Interestingly, the suggested timing of diversification is consistent with the hypothesis of a radiation that could have occurred contemporaneously with the diversification of their legume hosts.     

Traditional taxonomic groupings mask evolutionary history: a molecular phylogeny and new classification of the chromodorid nudibranchs

Chromodorid nudibranchs (16 genera, 300+ species) are beautiful, brightly colored sea slugs found primarily in tropical coral reef habitats and subtropical coastal waters. The chromodorids are the most speciose family of opisthobranchs and one of the most diverse heterobranch clades. Chromodorids have the potential to be a model group with which to study diversification, color pattern evolution, are important source organisms in natural products chemistry and represent a stunning and widely compelling example of marine biodiversity. Here, we present the most complete molecular phylogeny of the chromodorid nudibranchs to date, with a broad sample of 244 specimens (142 new), representing 157 (106 new) chromodorid species, four actinocylcid species and four additional dorid species utilizing two mitochondrial markers (16s and COI). We confirmed the monophyly of the Chromodorididae and its sister group relationship with the Actinocyclidae. We were also able to, for the first time, test generic monophyly by including more than one member of all 14 of the non-monotypic chromodorid genera. Every one of these 14 traditional chromodorid genera are either non-monophyletic, or render another genus paraphyletic. Additionally, both the monotypic genera Verconia and Diversidoris are nested within clades. Based on data shown here, there are three individual species and five clades limited to the eastern Pacific and Atlantic Oceans (or just one of these ocean regions), while the majority of chromodorid clades and species are strictly Indo-Pacific in distribution. We present a new classification of the chromodorid nudibranchs. We use molecular data to untangle evolutionary relationships and retain a historical connection to traditional systematics by using generic names attached to type species as clade names.     

Phylogenetic relationships in the Neotropical bruchid genus Acanthoscelides (Bruchinae, Bruchidae, Coleoptera)

Adaptation to host-plant defences through key innovations is a driving force of evolution in phytophagous insects. Species of the neotropical bruchid genus Acanthoscelides Schilsky are known to be associated with specific host plants. The speciation processes involved in such specialization pattern that have produced these specific associations may reflect radiations linked to particular kinds of host plants. By studying host-plant associations in closely related bruchid species, we have shown that adaptation to a particular host-plant (e.g. with a certain type of secondary compounds) could generally lead to a radiation of bruchid species at the level of terminal branches. However, in some cases of recent host shifts, there is no congruence between genetic proximity of bruchid species, and taxonomic similarity of host plants. At deeper branches in the phylogeny, vicariance or long-distance colonization events seem to be responsible for genetic divergence between well-marked clades rather than adaptation to host plants. Our study also suggests that the few species of Acanthoscelides described from the Old World, as well as Neotropical species feeding on Mimosoideae, are misclassified, and are more closely related to the sister genus Bruchidius .     

Ecological and evolutionary diversification of the seed beetle genus Stator (Coleoptera: Chrysomelidae: Bruchinae)

Ehrlich and Raven's (1964) hypothesis on coevolution has stimulated numerous phylogenetic studies that focus on the effects of plant defensive chemistry as the main ecological axis of phytophagous insect diversification. However, other ecological features affect host use and diet breadth and they may have very different consequences for insect evolution. In this paper, we present a phylogenetic study based on DNA sequences from mitochondrial and protein-coding genes of species in the seed beetle genus Stator, which collectively show considerable interspecific variation in host affiliation, diet breadth, and the dispersal stage of the seeds that they attack. We used comparative analyses to examine transitions in these three axes of resource use. We argue that these analyses show that diet breadth evolution is dependent upon colonizing novel hosts that are closely or distantly related to the ancestral host, and that oviposition substrate affects the evolution of host-plant affiliation, the evolution of dietary specialization, and the degree to which host plants are shared between species. The results of this study show that diversification is structured by interactions between different selective pressures and along multiple ecological axes.     

Phylogeny and host-plant association in the leaf beetle genus Trirhabda LeConte (Coleoptera: Chrysomelidae)

The leaf beetle genus Trirhabda contains 26 described species from the United States and Canada, feeding on host plants from the families Asteraceae and Hydrophyllaceae. In this study, we present a phylogeny for the genus that was reconstructed from mitochondrial COI and 12S rRNA fragments, nuclear ITS2 rRNA, and morphological characters. Both parsimony and mixed-model Bayesian likelihood analyses were performed. Under both methods, the mitochondrial and nuclear partitions support the same backbone phylogeny, as do the combined data. The utility of the molecular data is contrasted with the low phylogenetic signal among morphological characters. The phylogeny was used to trace the evolution of the host-plant association in Trirhabda. The recovered phylogeny shows that although the host-plant association is phylogenetically conservative, Trirhabda experienced one shift to a distantly related host-plant family, 6 shifts between host-plant tribes, and 6 between genera within tribes. The phylogeny reveals that Trirhabda were plesiomorphically adapted to tolerate complex secondary compounds of its host plants and this adaptation is retained in Trirhabda species, as evidenced by multiple shifts from chemically simpler host plants back to the more complex host plants.     

Host-plants shape insect diversity: Phylogeny, origin, and species diversity of native Hawaiian leafhoppers (Cicadellidae: Nesophrosyne)

Herbivorous insects and the plants on which they specialize, represent the most abundant terrestrial life on earth, yet their inter-specific interactions in promoting species diversification remains unclear. This study utilizes the discreet geologic attributes of Hawai'i and one of the most diverse endemic herbivore radiations, the leafhoppers (Hemiptera: Cicadellidae: Nesophrosyne), as a model system to understand the role of host-plant use in insect diversification. A comprehensive phylogeny is reconstructed to examine the origins, species diversification, and host-plant use of the native Hawaiian leafhoppers. Results support a monophyletic Nesophrosyne, originating from the Western Pacific basin, with a sister-group relationship to the genus Orosius. Nesophrosyne is characterized by high levels of endemicity according to individual islands, volcanoes, and geologic features. Clades demonstrate extensive morphologically cryptic diversity among allopatric species, utilizing widespread host-plant lineages. Nesophrosyne species are host-plant specific, demonstrating four dominant patterns of specialization that shape species diversification: (1) diversification through host switching; (2) specialization on widespread hosts with allopatric speciation; (3) repeated, independent shifts to the same hosts; and, (4) absence or low abundance on some host. Finally, evidence suggests competing herbivore radiations limit ecological opportunity for diversifying insect herbivores. Results provide evolutionary insights into the mechanisms that drive and shape this biodiversity.     

Intraclass Evolution and Classification of the Colpodea (Ciliophora)

ABSTRACT. Using nine new taxa and statistical inferences based on morphological and molecular data, we analyze the evolution within the class Colpodea. The molecular and cladistic analyses show four well‐supported clades: platyophryids, bursariomorphids, cyrtolophosidids, and colpodids. There is a widespread occurrence of homoplasies, affecting even conspicuous morphological characteristics, e.g. the inclusion of the micronucleus in the perinuclear space of the macronucleus. The most distinct changes in the morphological classification are the lack of a basal divergence into two subclasses and the split of the cyrtolophosidids into two main clades, differing mainly by the presence vs. absence of an oral cavity. The most complex clade is that of the colpodids. We partially reconcile the morphological and molecular data using evolutionary systematics, providing a scenario in which the colpodids evolved from a Bardeliella‐like ancestor and the genus Colpoda performed an intense adaptive radiation, giving rise to three main clades: Colpodina n. subord., Grossglockneriina, and Bryophryina. Three new taxa are established: Colpodina n. subord., Tillinidae n. fam., and Ottowphryidae n. fam. Colpodean evolution and classification are far from being understood because sequences are lacking for most species and half of their diversity is possibly undescribed.     

A phylogenetic perspective on 160 years of troubled taxonomy of Niphargus (Crustacea: Amphipoda)

Niphargus is the largest genus of freshwater amphipods. Its systematics from the species to the family level has always been problematic. This study is the first comprehensive phylogenetic treatment of the chiefly subterranean group in 160 years of its taxonomic history. It includes 103 niphargid species plus outgroups, representing about one‐third of all nominal species. The samples originated mainly from type localities or adjacent sites and covered most of the morphological variability of the genus. Character sampling included nuclear 28S and mitochondrial 12S rDNA sequences, and 122 morphological characters. Quantitative morphological traits were coded using two alternative methods. The first one searches for gaps in the variability range of each character, while the second one uses absolute differences between the standardized raw data as weights. Different data sets yielded alternative topologies. All data support the monophyly of Niphargidae, while Niphargopsis— another niphargid genus — was consistently nested within Niphargus, loosing justification for its separate status. We predict a similar fate for all or most of the remaining six small niphargid genera, which were not yet scrutinized phylogenetically. Different topologies agreed in species composition of five large, well‐supported clades, although the hierarchic relationships between them remain unresolved. These clades reject all previously proposed taxonomic subdivisions of Niphargus, implying a high degree of morphological homoplasy that renders any morphology‐based groups questionable. The clade members are distributed within well‐established zoogeographical regions that do not exceed 1300 km across the longest diagonal. These results provide a framework for future studies on niphargid systematics, the evolution of endemism and cryptic diversity in subterranean environments, the mechanisms leading to exceptional morphological heterogeneity, historical biogeography, and applied ecological issues.     

Defining the limits of taxonomic conservatism in host-plant use for phytophagous insects: molecular systematics and evolution of host-plant associations in the seed-beetle genus Bruchus Linnaeus (Coleoptera: Chrysomelidae: Bruchinae)

In this study, we have investigated the limits of taxonomic conservatism in host-plant use in the seed-beetle genus Bruchus. To reconstruct the insect phylogeny, parsimony and multiple partitioned Bayesian inference analyses were conducted on a combined data set of four genes. Permutation tests and both global and local maximum-likelihood optimizations of host preferences at distinct taxonomic levels revealed that host-fidelity is still discernible beyond the host-plant tribe level, suggesting the existence of more important than previously thought evolutionary constraints, which are further discussed in details. Our tree topologies are also mostly consistent with extant taxonomic groups. Through the analysis of this empirical data set we also provide meaningful insights on two methodological issues. First, Bayesian inference analyses suggest that partitioning by using codon positions greatly increase the accuracy of phylogenetical reconstructions. Regarding reconstruction of ancestral character states through maximum likelihood, the present study also highlights the usefulness of local optimizations. The issue of over-parameterization is also addressed, as the optimizations with the most parameter-rich models have returned the most counterintuitive results.     

Genetic divergence, speciation and biogeography of Mustelus (sharks) in the central Indo-Pacific and Australasia

The shark genus Mustelus is speciose, commercially important and systematically troublesome. We use a molecular approach combining inter and intra-specific data to investigate Mustelus species in the central Indo-Pacific and Australasia. Our analysis supports two Mustelus clades, one comprising species with no white spots and a placental reproductive mode and a second clade of white spotted, aplacental species. Levels of genetic divergence are low, especially among species in the white spotted, aplacental clade and this should be taken into account when employing molecular data to delineate species. Our data support the hypothesis of a radiation following dispersal from a northern hemisphere ancestor. Molecular dating suggests that localised speciation in Australasia may have occurred during the Pleistocene. We propose that some of the difficulties associated with Mustelus systematics relate to a recent radiation, particularly in the Australasian region.     

Phylogenetics and evolution of host-plant use in leaf-mining sawflies (Hymenoptera: Tenthredinidae: Heterarthrinae)

The habit of mining within leaves has evolved convergently in numerous plant-feeding insect taxa. Many leaf-mining groups contain a large number of species with distinct feeding preferences, which makes them highly suitable for studies on the evolutionary history of host-plant use and on the role of niche shifts in speciation. We aimed to clarify the origin, classification, and ecological evolution of the tenthredinid sawfly subfamily Heterarthrinae, which contains c. 150 leaf-mining species that collectively feed on over 20 plant genera around the world. For this, we reconstructed the phylogeny of representative heterarthrine species and diverse outgroups from the superfamily Tenthredinoidea on the basis of DNA sequence data collected from two mitochondrial (CoI and Cytb) and two nuclear (EF-1α and NaK) genes. Thereafter, we inferred the history of niche diversification within Heterarthrinae by plotting larval host-plant associations on the trees, and by contrasting a time-calibrated leaf-miner phylogeny with the phylogeny of their host plants. The results show that: (1) heterarthrine leaf-miners constitute a monophyletic group that arose from external-feeding blennocampine lineages within the Tenthredinidae c. 110-80 million years ago; (2) heterarthrines generally radiated well after their host taxa, and extant host-plant associations therefore result from a combination of host conservatism and occasional shifts among available plant taxa; and (3) diversification in Heterarthrinae apparently occurs by multiple mechanisms, including sympatric or allopatric ecological speciation, non-ecological allopatric speciation, and possibly allochronic speciation. Overall, both present and historical host-use patterns within the Heterarthrinae exhibit striking similarities to patterns found in co-occurring herbivore taxa.     

CONVERGENT AND CORRELATED EVOLUTION OF MAJOR LIFE‐HISTORY TRAITS IN THE ANGIOSPERM GENUS LEUCADENDRON (PROTEACEAE)

Natural selection is expected to cause convergence of life histories among taxa as well as correlated evolution of different life‐history traits. Here, we quantify the extent of convergence of five key life‐history traits (adult fire survival, seed storage, degree of sexual dimorphism, pollination mode, and seed‐dispersal mode) and test hypotheses about their correlated evolution in the genus Leucadendron (Proteaceae) from the fire‐prone South African fynbos. We reconstructed a new molecular phylogeny of this highly diverse genus that involves more taxa and molecular markers than previously. This reconstruction identifies new clades that were not detected by previous molecular study and morphological classifications. Using this new phylogeny and robust methods that account for phylogenetic uncertainty, we show that the five life‐history traits studied were labile during the evolutionary history of the genus. This diversity allowed us to tackle major questions about the correlated evolution of life‐history strategies. We found that species with longer seed‐dispersal distances tended to evolve lower pollen‐dispersal distance, that insect‐pollinated species evolved decreased sexual dimorphism, and that species with a persistent soil seed‐bank evolved toward reduced fire‐survival ability of adults.     

Phylogeny of the scathophagidae (Diptera, calyptratae) based on mitochondrial DNA sequences

The family Scathophagidae constitutes, together with members of the families Muscidae, Fannidae, and Anthomyiidae, the Muscoidea superfamily. The species Scathophaga stercoraria has been used extensively to investigate questions in animal ecology and evolution, particularly as a model system for studies of sperm competition and life history evolution. However, no phylogenetic studies have ever been performed on the Scathophagidae and the relationships within this family remain unclear. This study represents a molecular approach aimed at uncovering the phylogenetic relationships among 61 species representing 22 genera of Scathophagidae. A fragment of the terminal region of the mitochondrial gene COI (subunit I of the cytochrome oxidase gene) was sequenced in scathophagid species covering a wide geographic area, as well as a diverse spectrum of ecological habitats. Several clades grouping different genera and species have been identified, but the resolution power of the COI was insufficient to establish the exact relationships between these clades. The molecular data confirm the existence of a group consisting of the genera Delina, Chylizosoma, and Americina, which could represent the subfamily Delinae. Concerning the controversial position of the genus Phrosia, our data clearly suggest that it should be removed from the Delinae and placed within the genus Cordilura. Monophyly of most genera was confirmed, except for the genus Scathophaga, which should be divided into several different taxa.     

A cladistic analysis of Asarum (Aristolochiaceae) and implications for the evolution of herkogamy

A cladistic analysis of Asarum was conducted to examine relationships among species within the genus and to test the monophyly of several groups of taxa that have often been treated as segregate genera. Thirty-two species were drawn from throughout the range of the genus, representing a broad sample of sections and all segregate genera. The data matrix included 37 characters derived from various aspects of vegetative and floral morphology. A strict consensus of all most parsimonious trees suggests that Asarum s.l. is monophyletic and consists of two main clades: an Asarum clade, which is characterized by connate styles and inferior ovaries, and an Asiasarum-Hexastylis-Heterotropa clade, which is characterized by ridges on the inner perianth surface, dorsal stigmas, and bifid style extensions. The latter is a large and morphologically diverse clade that includes the North American segregate Hexastylis and two Asiatic segregates. Examination of pollination mechanisms in the context of this phylogeny supports the conclusion that herkogamy, and thus obligate insect pollination, is derived from a plesiomorphic condition of autonomous self-pollination. Associated with herkogamy are characters such as glandular trichomes and other ornamentation of the surface of the calyx that probably represent increased specialization to attract insect pollinators. This study also indicates that chromosomal evolution has occurred via aneuploid decrease from an ancestral chromosome number of 2n = 26 to 2n = 24 in Heterotropa. The recognition of two subgenera, subgenus Asarum and subgenus Heterotropa, corresponding to the two clades in the cladistic analysis, is recommended.     

Insights on speciation patterns in the genus Iberochondrostoma (Cyprinidae): evidence from mitochondrial and nuclear data

In this paper, the patterns of cladogenesis in the cyprinid fish genus Iberochondrostoma were analysed using a mitochondrial (cytochrome b) and a nuclear (beta-actin) gene fragment. The two genes yielded discordant results. While the cytochrome b gene yielded a fully dichotomous tree, where all species of the genus are monophyletic, the much slower beta-actin gene yielded star-like relationships. However, when information from both genes was considered together, the data suggested the persistence of a very large central unit from which at least two peripheral clades arose at different times. This pattern which is akin to peripatric speciation was shown to be compatible with the paleogeographical information available. It is suggested that combining the techniques of phylogeny and phylogeography and the use of multiple markers varying in their rate of evolution may enrich our understanding of speciation and evolution of clades beyond species level.