18S rRNA secondary structure and phylogenetic position of Peloridiidae (Insecta, hemiptera)

A secondary structure model for 18S rRNA of peloridiids, relict insects with a present-day circumantarctic distribution, is constructed using comparative sequence analysis, thermodynamic folding, a consensus method using 18S rRNA models of other taxa, and support of helices based on compensatory substitutions. Results show that probable in vivo configuration of 18S rRNA is not predictable using current free-energy models to fold the entire molecule concurrently. This suggests that refinements in free-energy minimization algorithms are needed. Molecular phylogenetic datasets were created using 18S rRNA nucleotide alignments produced by CLUSTAL and rigorous interpretation of homologous position based on certain secondary substructures. Phylogenetic analysis of a hemipteran data matrix of 18S rDNA sequences placed peloridiids sister to Heteroptera. Resolution of affiliations between the three main euhemipteran lineages was unresolved. The peloridiid 18S RNA model presented here provides the most accurate template to date for aligning homologous nucleotides of hemipteran taxa. Using folded 18S rRNA to infer homology of character as morpho-molecular structures or nucleotides and scoring particular sites or substructures is discussed.     

Novelty in phylogeny of gastrotricha: evidence from 18S rRNA gene

Gastrotricha form a phylum which is crucial for defining the origin of pseudocoelomates, in that they share a number of characters with Rotifera and Nematoda but also with acoelomates, and even the evolutionary relationships within the phylum are anything but defined. For this reason the first extensive molecular data on Gastrotricha from the 18S rRNA sequences of both orders have been obtained and analyzed. Sequence analyses show that the phylum Gastrotricha is strictly monophyletic along an evolutionary line quite distinct from that of both Rotifera and Nematoda. A new view of the evolutionary history of the phylum Gastrotricha is put forward, in which Chaetonotida, and not Macrodasyida, are the most primitive forms of the group, contrary to the commonly held view. A polyphyletic origin of aschelminthes is supported, and the misleading term pseudocoelomates should be discarded.     

A preliminary phylogeny of the Pentatomomorpha (Hemiptera: Heteroptera) based on nuclear 18S rDNA and mitochondrial DNA sequences

Pentatomomorpha is the second suborder in size only to Cimicomorpha in Heteroptera. However, the phylogenetic relationships among members of the suborder are not well established. Sequences from partial nuclear ribosomal 18S gene and mitochondrial COX1 gene were analyzed separately and in combination to generate a preliminary molecular phylogeny of Pentatomomorpha based on 40 species representing 17 putative families. Analyses of the combined sequence data provided a better-resolved and more robust hypothesis of Pentatomomorpha phylogeny than did separate analyses of the individual genes. The phylogenies were mostly congruent with morphological studies. Results strongly supported the monophyly of the infraorder Pentatomomorpha, and the placement of Aradoidea as sister to Trichophora. The monophyletic Trichophora was grouped into two major lineages, one being the superfamily Pentatomoidea, and the other comprising Lygaeoidea, Coreoidea, and Pyrrhocoroidea. The analysis of the ML and ME trees of combined dataset supported the monophyletic Pentatomoidea. In all analysis the Pyrrhocoroidea was polyphyletic; the monophyletic Lygaeoidea was supported only in the analysis of ME tree, and Coreoidea was polyphyletic except in the MP tree of combined dataset. The molecular and morphylogical data both indicated that the family Coreoidae should be revised subsequently. Our phylogenetic results suggested that the COX1 segment alone might not be an optimal molecular marker for the phylogeny of Pentatomomorpha.     

A molecular phylogeny of heterodont bivalves (Mollusca: Bivalvia: Heterodonta): new analyses of 18S and 28S rRNA genes

A new molecular phylogeny is presented for the highly diverse, bivalve molluscan subclass Heterodonta. The study, the most comprehensive for heterodonts to date, used new sequences of 18S and 28S rRNA genes for 103 species from 49 family groups with species of Palaeoheterodonta (Trigoniidae, Margaritiferidae and Unionidae) as outgroups. Results confirm previous analyses that the Carditidae/Astartidae/Crassatellidae clade is basal to all other heterodonts including Anomalodesmata (often classified as a separate subclass or order). Thyasiroidea occupy a near basal position between the Crassatelloidea and Anomalodesmata. Lucinidae form a well‐supported monophyletic group distinct from Thyasiridae and Ungulinidae. The Solenoidea and Hiatelloidea link as sister groups distant from the Tellinoidea and Myoidea, respectively, where they had been previously associated. The position of the Gastrochaenidae is unstable but does not group with myoidean taxa. Species of four families of Galeommatoidea form a clade that also includes Sportellidae of the Cyamioidea. The Cardioidea and Tellinoidea form highly supported, long branched, individual clades but group as sister taxa. A major clade including Veneroidea, Mactroidea, Myoidea and other families is given the unranked name Neoheterodontei. There is no support for a separate order Myoida (Myoidea and Pholadoidea). Dreissenidae group within the clade including Myidae, Corbulidae, Pholadidae and Teredinidae. The Corbiculoidea is confirmed as polyphyletic with the Sphaeriidae and Corbiculidae forming separate clades within the Neoheterodontei; Corbiculidae grouping with the Glauconomidae. Hemidonacidae are unrelated to the Cardiidae, as previously proposed, but nest within the Neoheterodontei. The Gaimardiidae group near to the Ungulinidae and not with Cyamioidea where most recently classified. The family Ungulinidae, previously classified in the Lucinoidea, forms a well‐supported clade within the Neoheterodontei and is elevated to superfamily rank — Ungulinoidea. The monophyletic status of Glossoidea, Arcticoidea and Veneroidea is unconfirmed. A brief review of the fossil record of the heterodonts indicates that the basal clades of Crassatelloidea, Anomalodesmata and Lucinoidea diverged very early in the Lower Palaeozoic. Other groups such as the Hiatelloidea, Solenoidea, Gastrochaenidae probably were of late Palaeozoic origins. The Cardioidea and Tellinoidea originated in the Triassic while major groups of Neoheterodontei radiated in the Late Mesozoic. The phylogenetic position of the Thyasiroidea and Galeommatoidea suggests a longer fossil history than has so far been recognized.     

New insight into the phylogeny of Mesozoa: Evidence from the 18S and 28S rRNA genes

The phylogenetic relationships of two Mesozoa groups were studied by the comparative analysis of complete sequences of 18S and 28S rRNA. Two groups of Mesozoa were found to form a statistically supported clade in phylogenetic trees. The results of the analysis placed Mesozoa in the Spiralia group of Lophotrochozoa and showed the tendency of Mesozoa to converge with one of the Annelida groups.     

Evolution of the planthoppers (Insecta: Hemiptera: Fulgoroidea)

The planthopper superfamily Fulgoroidea (Insecta: Hemiptera) comprises approximately 20 described insect families, depending on which classification is followed. Multiple competing hypotheses of fulgoroid phylogeny have been published, based on either morphological character coding or DNA sequence data; however, those hypotheses disagree in several key aspects regarding the evolution of planthoppers. The current paper seeks to test these hypotheses, including the Asche (Asche, M. 1987. Preliminary thoughts on the phylogeny of Fulgoromorpha (Homoptera Auchenorrhyncha). In: Proceedings of the 6th Auchenorrhyncha Meeting, Turin, Italy, 7-11 September, 1987, pp. 47-53.) hypothesis of a trend in ovipositor structure, which may be correlated with planthopper feeding ecology. Presented here are phylogenetic reconstructions of Fulgoroidea based on analysis of DNA nucleotide sequence data from four loci (18S rDNA, 28S rDNA, Histone 3, and Wingless) sequenced from 83 exemplar taxa representing 18 planthopper families and outgroups. Data sets were analyzed separately and in various combinations under the maximum parsimony criterion, and the total combined dataset was analyzed via both maximum parsimony and partitioned Bayesian criteria; results of the combined analyses were concordant across reconstruction paradigms. Relationships recovered suggest several major planthopper lineages, including: (1) Delphacidae+Cixiidae; (2) Kinnaridae+Meenoplidae; (3) Fulgoridae+Dictyopharidae; (4) Lophopidae+Eurybrachidae (possibly+Flatidae); (5) Ricaniidae+Caliscelidae (possibly+Tropiduchidae). Results also suggest the placement of Achilixiidae outside of Cixiidae and of Tettigometridae as one of the more recently diversified lineages within Fulgoroidea. The resulting phylogeny supports Asche's (1987) hypothesis of a functional trend in ovipositor structure across families.     

Higher level phylogeny and the first divergence time estimation of Heteroptera (Insecta: Hemiptera) based on multiple genes

Heteroptera, or true bugs, are the largest, morphologically diverse and economically important group of insects with incomplete metamorphosis. However, the phylogenetic relationships within Heteroptera are still in dispute and most of the previous studies were based on morphological characters or with single gene (partial or whole 18S rDNA). Besides, so far, divergence time estimates for Heteroptera totally rely on the fossil record, while no studies have been performed on molecular divergence rates. Here, for the first time, we used maximum parsimony (MP), maximum likelihood (ML) and Bayesian inference (BI) with multiple genes (18S rDNA, 28S rDNA, 16S rDNA and COI) to estimate phylogenetic relationships among the infraorders, and meanwhile, the Penalized Likelihood (r8s) and Bayesian (BEAST) molecular dating methods were employed to estimate divergence time of higher taxa of this suborder. Major results of the present study included: Nepomorpha was placed as the most basal clade in all six trees (MP trees, ML trees and Bayesian trees of nuclear gene data and four-gene combined data, respectively) with full support values. The sister-group relationship of Cimicomorpha and Pentatomomorpha was also strongly supported. Nepomorpha originated in early Triassic and the other six infraorders originated in a very short period of time in middle Triassic. Cimicomorpha and Pentatomomorpha underwent a radiation at family level in Cretaceous, paralleling the proliferation of the flowering plants. Our results indicated that the higher-group radiations within hemimetabolous Heteroptera were simultaneously with those of holometabolous Coleoptera and Diptera which took place in the Triassic. While the aquatic habitat was colonized by Nepomorpha already in the Triassic, the Gerromorpha independently adapted to the semi-aquatic habitat in the Early Jurassic.     

Entomologically famous, evolutionarily unexplored: The first phylogeny of the lanternfly family Fulgoridae (Insecta: Hemiptera: Fulgoroidea)

Lanternflies (Insecta: Hemiptera: Fulgoridae) are frequently used as examples of unusual morphological evolution, with some species (such as the peanut-headed bug, Fulgora laternaria Linnaeus) also ubiquitously cited as icons of tropical insect biodiversity. Despite that entomological notoriety, the phylogeny of this charismatic planthopper family has never before been studied. Presented here are the results of a phylogenetic investigation of Fulgoridae based on DNA nucleotide sequence data from five genetic loci (18S rDNA, 28S rDNA, histone 3, wingless, and cytochrome oxidase I). The resulting topologies are used to test the higher classification of Fulgoridae, which is based primarily on characters associated with the curious head morphology of many included species. Analyses include a taxonomic sample of 69 fulgorid species representing 46 (of 110) genera, 10 (of 11) tribes, and all 8 currently recognized subfamilies. Results of this study: (1) demonstrate the need for a revised classification of Fulgoridae, particularly at the higher taxonomic levels; (2) suggest that the genus Zanna is excluded from a monophyletic Fulgoridae; (3) indicate that there have been multiple losses of the extended head process across fulgorid evolution, with what appears to be convergence (in shape and/or loss) in distantly related lineages; and (4) suggest two alternative biogeographic hypotheses to explain the distribution of extant Fulgoridae, with either an Old World origin and a single subsequent colonization of the New World, or a contemporaneous diversification of Old and New World lineages.     

On the constitution and phylogeny of Staphyliniformia (Insecta: Coleoptera)

The Staphyliniformia is one of the most diverse lineages of Coleoptera, with representatives occupying every conceivable non-marine niche. Phylogenetic relationships among its varied families and lower taxa have defied resolution. The problem has been further complicated by the recent suggestion that another major coleopteran series, Scarabaeiformia, is derived from within it. Here we present the first phylogenetic analyses, based on 18S rDNA sequences and morphological data, to explicitly examine this possibility. Thorough evaluation of alternative alignments and tree construction methods support the contention that Scarabaeiformia is derived from within Staphyliniformia. Though the analyses yielded strong support for few family level groupings within the expanded Staphyliniformia, they conclusively support a close relationship between Hydraenidae and Ptiliidae, which has often been debated. The primary factor hindering additional resolution appears to be the inconsistent rate of divergence in 18S among these taxa.     

Influence of data conflict and molecular phylogeny of major clades in Cimicomorphan true bugs (Insecta: Hemiptera: Heteroptera)

Cimicomorpha, which consists of 16 families representing more than 19,400 species, is the largest infraorder in Heteroptera, Insecta. We present the first molecular phylogenetic investigation of family relationships of Cimicomorpha, including 46 taxa from 12 of 16 Cimicomorphan families. Three genes, with a total of 3277 bp of sequence data (nuclear 18S rDNA: 2022 bp, 28S rDNA: 755 bp, and mitochondrial 16S rDNA: 498 bp) were analyzed. Data partitions were analyzed separately and in combination, by employing ML (maximum likelihood), MP (maximum parsimony), and Bayesian methods. As saturation was detected in substitutions of 16S rDNA, influence of data conflict in combined analyses was further explored by three methods: the incongruence length difference (ILD) test, the partitioned Bremer support (PBS), and the partition addition bootstrap alteration approach (PABA). PBS and PABA approaches suggested that 16S rDNA was not very suitable for addressing relationships at this level in Cimicomorpha. Our results also supported the nabid-cimicoid lineage for Cimicoidea proposed by Schuh and Stys [Schuh, R.T., Stys, P., 1991. Phylogenetic analysis of Cimicomorphan family relationships (Heteroptera). J. NY Entomol. Soc. 99 (3), 298-350]. Data incongruence and the utility of the three genes were briefly discussed.     

Predicted Secondary Structure for 28S and 18S rRNA from Ichneumonoidea (Insecta: Hymenoptera: Apocrita): Impact on Sequence Alignment and Phylogeny Estimation

We utilize the secondary structural properties of the 28S rRNA D2–D10 expansion segments to hypothesize a multiple sequence alignment for major lineages of the hymenopteran superfamily Ichneumonoidea (Braconidae, Ichneumonidae). The alignment consists of 290 sequences (originally analyzed in Belshaw and Quicke, Syst Biol 51:450–477, 2002) and provides the first global alignment template for this diverse group of insects. Predicted structures for these expansion segments as well as for over half of the 18S rRNA are given, with highly variable regions characterized and isolated within conserved structures. We demonstrate several pitfalls of optimization alignment and illustrate how these are potentially addressed with structure-based alignments. Our global alignment is presented online at (http://hymenoptera.tamu.edu/rna) with summary statistics, such as basepair frequency tables, along with novel tools for parsing structure-based alignments into input files for most commonly used phylogenetic software. These resources will be valuable for hymenopteran systematists, as well as researchers utilizing rRNA sequences for phylogeny estimation in any taxon. We explore the phylogenetic utility of our structure-based alignment by examining a subset of the data under a variety of optimality criteria using results from Belshaw and Quicke (2002) as a benchmark.Access to on-line data: http://hymenoptera.tamu.edu/rna; username, ichs; password, ichzzz     

蜡蝉总科(昆虫纲:半翅目)系统分类研究进展

蜡蝉总科隶属于半翅目头喙亚目,大致可分为18～21个科。中国记载16科。蜡蝉总科的科级系统发育关系一直存在着广泛争议,至今尚无一个为大多数学者所接受的分类系统。因此,研究蜡蝉总科的系统发育具有重要的理论价值。本文从种及种下阶元的分类鉴定、种上阶元的系统发育、生物地理学等方面对蜡蝉总科的分类研究进行综述。     

A contribution to sedentary polychaete phylogeny using 18S rRNA sequence data

The phylogenetic position of Annelida as well as its ingroup relationships are a matter of ongoing debate. A molecular phylogenetic study of sedentary polychaete relationships was conducted based on 70 sequences of 18S rRNA, including unpublished sequences of 18 polychaete species. The data set was analysed with maximum parsimony and maximum likelihood methods. Clade robustness was estimated by parsimony-bootstrapping and jackknifing, decay index, and clade support, as well as a posteriori probability tests using Bayesian inference. Irrespective of the applied method, some traditional sedentary polychaete taxa, such as Cirratulidae, Opheliidae, Orbiniidae, Siboglinidae and Spionidae, were recovered by our phylogenetic reconstruction. A close relationship between Orbiniidae and Questa received a particularly strong support. Echiura appears to be a polychaete ingroup taxon which is closely related to Dasybranchus (Capitellidae). As in previous molecular analyses, no support was found for the monophyly of Annelida nor for that of Polychaeta. However, we suggest that an increase in taxon sampling may yield additional resolution in the reconstruction of polychaete ingroup phylogeny, although the difficulties in reconstructing the basal phylogenetic relationships within Annelida may be due to their rapid radiation.     

Aligned 18S for Zoraptera (Insecta): phylogenetic position and molecular evolution

The order Zoraptera (angel insects) is one of the least known insect groups, containing only 32 extant species. The phylogenetic position of Zoraptera is poorly understood, but it is generally thought to be closely related to either Paraneoptera (hemipteroid orders: booklice, lice, thrips, and bugs), Dictyoptera (blattoid orders: cockroaches, termites, and mantis), or Embioptera (web spinners). We inferred the phylogenetic position of Zoraptera by analyzing nuclear 18S rDNA sequences, which we aligned according to a secondary structure model. Maximum likelihood and Bayesian analyses both supported a close relationship between Zoraptera and Dictyoptera with relatively high posterior probability. The 18S sequences of Zoraptera exhibited several unusual properties: (1) a dramatically increased substitution rate, which resulted in very long branches; (2) long insertions at helix E23; and (3) modifications of secondary structures at helices 12 and 18.     

An 18S rDNA-based molecular phylogeny of aphidiinae (Hymenoptera: braconidae)

We have obtained a molecular phylogeny of the subfamily Aphidiinae (Hymenoptera: Braconidae) by sequencing the 18S rDNA in 37 aphidiine taxa. Approximately 1857 nucleotides were sequenced in each species. Evolutionary relationships were established by comparing the results of maximum-parsimony, maximum-likelihood, and distance analyses. The most variable region of this gene, V4 (approx 403 nucleotides), was employed to establish the basality of the tribe Ephedrini within this subfamily. All phylogenetic reconstructions yielded trees with very similar topologies that confirmed the existence of two of the four traditionally accepted tribes, Ephedrini and Praini, but questioned the existence of Trioxini and Aphidiini. To better ascertain the status of some groups, the same analyses were repeated with a reduced taxonomic sample in which some species that produced systematic errors in the former phylogenetic reconstructions had been removed. The results from this second analysis favor either the three-tribes hypothesis (Ephedrini, Praini, and Aphidiini) or a new classification with at least five tribes (Ephedrini, Praini, Monoctonini, Trioxini, and Aphidiini). The 18S rDNA gene is a useful marker to recover relationships not only at the tribe but also at the subtribe and genus levels in this group. The natural status of some traditionally accepted clusters is also corroborated with the present data whereas the placement of other clusters is questioned or remains unresolved.     

Biogeography of the wax scales (Insecta: Hemiptera: Coccidae: Ceroplastinae)

The modern and native distributions of wax scales are documented and an area cladogram including seventy species is presented. Wax scales are distributed worldwide, but most species are native to either South America or Africa. Their native distribution pattern is discussed in relation to their host-plant specificity and to the dispersal and vicariance theories of biogeography. The vicariance theory is preferred, because the pattern can be explained satisfactorily by plate tectonics but not by dispersal from a centre of origin. The wax scale group probably originated in the combined South American–African continent at least 97 million yr ago.     

Revision of New Zealand Orthotylinae (Insecta: Hemiptera: Miridae)

Abstract

Six genera and 11 species of Orthotylinae are now known from New Zealand. Zanchius Distant is newly recorded. One new genus, Tridiplous, and seven new species of endemic New Zealand Orthotylini are described and illustrated. Four new species are described in Tridiplous: T. burrus, T. parvapiatus, T. penmani, and T. virens, and three new species are described in Zanchius: Z. ater, Z. rubicrux, and Z. totus. Keys are provided to identify New Zealand taxa of the tribes of Orthotylinae, genera and species of Halticini, genera of Orthotylini, and species of Tridiplous and Zanchius. The bugs are illustrated with colour habitus photos, and drawings including male and female genitalia. Economic importance is discussed.     

Phylogeny of the major lineages of Membracoidea (Insecta: Hemiptera: Cicadomorpha) based on 28S rDNA sequences

Analysis of sequences from a 3.5-kb region of the nuclear ribosomal 28S DNA gene spanning divergent domains D2-D10 supports the hypothesis, based on fossil, biogeographic, and behavioral evidence, that treehoppers (Aetalionidae and Membracidae) are derived from leafhoppers (Cicadellidae). Maximum-parsimony analysis indicated that treehoppers are the sister group of a lineage comprising the currently recognized cicadellid subfamilies Agalliinae, Megophthalminae, Adelungiinae, and Ulopinae. Based on this phylogenetic estimate, the derivation of treehoppers approximately coincided with shifts in physiology and behavior, including loss of brochosome production and a reversal from active, jumping nymphs to sessile, nonjumping nymphs. Myerslopiidae, traditionally placed as a tribe of the cicadellid subfamily Ulopinae, represented a basal lineage distinct from other extant membracoids. The analysis recovered a large leafhopper lineage comprising a polyphyletic Deltocephalinae (sensu stricto) and its apparent derivatives Koebeliinae, Eupelicinae (polyphyletic), Selenocephalinae, and Penthimiinae. Clades comprising Macropsinae, Neocoelidiinae, Scarinae, Iassinae, Coelidiinae, Eurymelinae + Idiocerinae, Evacanthini + Pagaroniini, Aphrodinae + Ledrinae (in part), Stenocotini + Tartessinae, and Cicadellini + Proconiini were also recovered with moderate to high branch support. Cicadellinae (sensu lato), Ledrinae, Typhlocybinae, and Xestocephalinae were consistently polyphyletic on the most-parsimonious topologies, but constraining these groups to be monophyletic did not significantly increase the length of the cladograms. Relationships among the major lineages received low branch support, suggesting that more data are needed to provide a robust phylogenetic estimate.     

Performance of 18S rRNA in Littorinid Phylogeny (Gastropoda: Caenogastropoda)

In the past, 18S rRNA sequences have proved to be very useful for tracing ancient divergences but were rarely used for resolving more recent ones. Moreover, it was suggested that the molecule does not contain useful information to resolve divergences which took place during less than 40 Myr. The present paper takes littorinid phylogeny as a case study to reevaluate the utility of the molecule for resolving recent divergences. Two data sets for nine species of the snail family Littorinidae were analyzed, both separately and combined. One data set comprised 7 new complete 18S rRNA sequences aligned with 2 published littorinid sequences; the other comprised 12 morphological, 1 biochemical, and 2 18S rRNA secondary structure characters. On the basis of its ability to confirm generally accepted relationships and the congruence of results derived from the different data sets, it is concluded that 18S rRNA sequences do contain information to resolve ``rapid' cladogenetic events, provided that they occurred in the not too distant past. 18S rRNA sequences yielded support for (1) the branching order (L. littorea, (L. obtusata, (L. saxatilis, L. compressa))) and (2) the basal position of L. striata in the Littorina clade. Received: 6 February 1998 / Accepted: 20 March 1998