Revisiting habitat and lifestyle transitions in Heteroptera (Insecta: Hemiptera): insights from a combined morphological and molecular phylogeny.

Heteroptera, the true bugs, are part of the largest clade of non-holometabolous insects, the Hemiptera, and include > 42 000 described species in about 90 families. Despite progress in resolving phylogenetic relationships between and within infraorders since the first combined morphological and molecular analysis published in 1993 (29 taxa, 669 bp, 31 morphological characters), recent hypotheses have relied entirely on molecular data. Weakly supported nodes along the backbone of Heteroptera made these published phylogenies unsuitable for investigations into the evolution of habitats and lifestyles across true bugs. Here we present the first combined morphological and molecular analyses of Heteroptera since 1993, using 135 taxa in 60 families, 4018 aligned bp of ribosomal DNA and 81 morphological characters, and various analytical approaches. The sister-group relationship of the predominantly aquatic Nepomorpha with all remaining Heteroptera is supported in all analyses, and a clade formed by Enicocephalomorpha, Dipsocoromorpha and Gerromorpha in some. All analyses recover Leptopodomorpha + (Cimicomorpha + Pentatomomorpha), mostly with high support. Parsimony- and likelihood-based ancestral state reconstructions of habitats and lifestyles on the combined likelihood phylogeny provide new insights into the evolution of true bugs. The results indicate that aquatic and semi-aquatic true bugs invaded these habitats three times independently from terrestrial habitats in contrast to a recent hypothesis. They further suggest that the most recent common ancestor of Heteroptera was predacious, and that the two large predominantly phytophagous clades (Trichophora and Miroidea) are likely to have derived independently from predatory ancestors. We conclude that by combining morphological and molecular data and employing various analytical methods our analyses have converged on a relatively well-supported hypothesis of heteropteran infraordinal relationships that now requires further testing using phylogenomic and more extensive morphological datasets.

Areas of endemism in the Nearctic: a case study of 1339 species of Miridae (Insecta: Hemiptera) and their plant hosts.

Areas of endemism are essential first hypotheses in investigating historical biogeography, but there is a surprising paucity of such hypotheses for the Nearctic region. Miridae, the plant bugs, are an excellent taxon to study in this context, because this group combines high species diversity, often small distribution ranges, a history of modern taxonomic revisions, and comprehensive electronic data capture and data cleaning that have resulted in an exceptionally error-free geospatial data set. Many Miridae are phytophagous and feed on only one or a small number of host plant species. The programs ndm/vndm are here used on plant bug and plant data sets to address two main objectives: (i) identify areas of endemism for plant bugs based on parameters used in a recent study that focused on Nearctic mammals; and (ii) discuss hypotheses on areas of endemism based on plant bug distributions in the context of areas identified by their host plant species. Given the narrow distribution ranges of many species of Miridae, the analytical results allow for tests of the prediction that areas of endemism for Miridae are smaller and more numerous, especially in the Western Nearctic, than are those of their host plants. Analyses of the default plant bug data set resulted in 45 areas of endemism, 35 of them north of Mexico and many located in the Western Nearctic; areas in the Nearctic are more numerous and smaller than those identified by mammals. The host plant data set resulted in ten areas of endemism, and even though the size range of areas is similar between the Miridae and plant data sets, the average area size is smaller in the Miridae data set. These results allow for the conclusion that the Miridae indeed present a valuable model system to investigate areas of endemism in the Nearctic.

Total-evidence phylogenetic analysis and reclassification of the Phylinae (Insecta: Heteroptera: Miridae), with the recognition of new tribes and subtribes and a redefinition of Phylini.

The subfamily Phylinae (Heteroptera: Miridae) is one of the largest subfamilies of plant bugs and in the most recent classification comprised six tribes: Pilophorini, Hallodapini, Auricillocorini, Phylini, Pronotocrepini, and Leucophoropterini. Phylogenetic analyses of the subfamily using dynamic homology (POY), parsimony (TNT), and model-based (RAxML) methods are presented. A dataset comprising both morphological and molecular characters (16S, 18S, 28S, and COI-COII) was assembled for taxon samples of 164 ingroup and nine outgroup taxa. A reclassification of the subfamily based on the POY analysis is presented, recognizing nine tribes and nine subtribes. The Auricillocorini is synonymized with the Hallodapini and the Pronotocrepini with the Cremnorrhini; the Phylini was found to be polyphyletic and is redefined; the Semiini and Nasocorini are resurrected and redefined; and the Decomiini and Coatonocapsiniare presented as new tribes. The Hallodapini, rather than the Pilophorini, was found to be the sister-group to the remaining Phylinae.

Systematics and evolution of Heteroptera: 25 years of progress.

Heteroptera, or true bugs, are part of the most successful radiation of nonholometabolous insects. Twenty-five years after the first review on the influence of cladistics on systematic research in Heteroptera, we summarize progress, problems, and future directions in the field. The few hypotheses on infraordinal relationships conflict on crucial points. Understanding relationships within Gerromorpha, Nepomorpha, Leptopodomorpha, Cimicomorpha, and Pentatomomorpha is improving, but progress within Enicocephalomorpha and Dipsocoromorpha is lagging behind. Nonetheless, the classifications of several superfamily-level taxa within the Pentatomomorpha, such as Aradoidea, Coreoidea, and Pyrrhocoroidea, are still unaffected by cladistic studies. Progress in comparative morphology is slow and drastically impedes our understanding of the evolution of major clades. Molecular systematics has dramatically contributed to accelerating the generation and testing of hypotheses. Given the fascinating natural history of true bugs and their status as model organisms for evolutionary studies, integration of cladistic analyses in a broader biogeographic and evolutionary context deserves increased attention.

emGraziaphylus/em, a new genus and two new species of emEremophila/em-feeding br /(Scrophulariceae) Phylinae (Insecta: Heteroptera: Miridae) from Australia.

The new genus Graziaphylus is described to include two new species, Graziaphylus joceliae and G. nigripes. This new taxon is part of a large suite of Heteroptera from Australia known to feed on species of Eremophila (Scrophulariaceae), including other members of the Miridae as well as the Tingidae. The appearance of Graziaphylus spp. is similar to that of Xiphoidellus Weirauch and Schuh from Australia, and to Chlamydatus Fieber from the Northern Hemisphere (among other genera), but with scalelike setae, and elongate pulvilli adnate over most of the claw length, and the distinctive structure of the male genitalia.

Systematic methods, fossils, and relationships within Heteroptera (Insecta).

Three recent papers dealing with phylogenetic relationships within the Heteroptera are discussed and analysed. A character set representing 43 taxa and 78 characters is used to test theories presented in those papers. The conclusions of Grimaldi and Engel concerning the placement of the Cretaceous fossil taxon Cretopiesma in the Piesmatidae are rejected in favour of placement in the Aradidae. The placement by Nel et al. of Protodoris from Eocene amber of the Paris Basin in the Thaumastocoridae is considered ambiguous because it has none of the diagnostic characters of that family group and is therefore regarded as incertae sedis. The arguments of Sweet concerning the elevation of the Aradoidea to infraordinal status on the basis of autapomorphies are also treated as invalid. General arguments against the use of phenetic methods in palaeontology, and ad hoc approaches under the guise of cladistics, are offered, with the conclusion that rigorous cladistic analyses are a prerequisite to testable conclusions concerning the placement of fossil and Recent taxa. © The Willi Hennig Society 2009.

Phylogenetic relationships of family groups in Pentatomoidea based on morphology and DNA sequences (Insecta: Heteroptera).

Phylogenetic relationships within the Pentatomoidea are investigated through the coding and analysis of character data derived from morphology and DNA sequences. In total, 135 terminal taxa were investigated, representing most of the major family groups; 84 ingroup taxa are coded for 57 characters in a morphological matrix. As many as 3500 bp of DNA data are adduced for each of 52 terminal taxa, including 44 ingroup taxa, comprising the 18S rRNA, 16S rRNA, 28S rRNA, and COI gene regions. Character data are analysed separately and in the form of a total evidence analysis. Major conclusions of the phylogenetic analysis include: the concept of Urostylididae is restricted to that of earlier authors; the Saileriolinae is raised to family rank and treated as the sister group of all Pentatomoidea exclusive of Urostylididae sensu stricto; a broadly conceived Cydnidae, as recognized by Dolling, 1981, is not supported; the placement of Thaumastellidae within the Pentatomoidea is affirmed and the taxon is recognized at family rank rather than as a subfamily of Cydnidae, although its exact phylogenetic position within the Pentatomoidea remains equivocal; the Parastrachiinae is treated as also including Dismegistus Amyot & Serville and placed within a broadly conceived Corimelaenidae, the latter group being treated at family rank; the family-group taxa Dinidoridae and Tessaratomidae probably represent a monophyletic group, but the recognition of monophyletic subgroups will benefit from additional representation in the sequence data set; and the Lestoniidae is treated as the sister group of the Acanthosomatidae. The Acanthosomatidae and Scutelleridae are consistently recovered as monophyletic. The monophyly of the Pentatomidae appears unequivocal, inclusive of the Aphylinae and Cyrtocorinae, on the basis of morphology, the latter two taxa not being represented in the molecular data set. © The Willi Hennig Society 2008.

Macrotylushenryi, a new species of Pelargonium-feeding Cremnorrhinina from South Africa (Hemiptera, Miridae, Phylinae, Cremnorrhinini).

Macrotylushenryi is described as a new species from South Africa. This new taxon is recorded as feeding on species of Pelargonium (Geraniaceae) in the Western Cape. Documentation is provided in the form of diagnosis, description, habitus photographs, scanning electron micrographs, illustrations and images of genitalic structures, detailed distributional data, host plant information, and images of hosts and habitats. Morphological traits are similar to species of Macrotylus Fieber from the Northern Hemisphere, but coloration is substantially variable, and the structure of the male genitalia is distinctive.

Henryognathusthomasi, a new genus and new species of Arctostaphylos-feeding plant bug from western North America (Miridae, Phylinae, Phylini).

Henryognathus, new genus, with the single included new species H.thomasi, is described from western North America. The taxon is recorded as feeding on species of Arctostaphylos (Ericaceae) in California and Arizona. Coloration and many morphological attributes are similar to species of Plagiognathus Fieber, but the structure of the male genitalia is distinctive.

PHYLOGENETIC, HOST AND BIOGEOGRAPHIC ANALYSES OF THE PILOPHORINI (HETEROPTERA: MIRIDAE: PHYLINAE).

A cladistic analysis is performed for 114 species of Pilophorini, using 73 characters with 190 states. Illustrations or figure references to the literature are provided for most characters. The resultant Nelson (strict) consensus cladogram is used as a justification for the recognition of the following genera: Alepidiella Poppius, Aloea Linnavuori, Druthmarus Distant, Hypseloecus Reuter, Neoambonea Schuh, Parambonea Schuh, Parasthenaridea Miller, Pherolepis Kulik (paraphyletic), Pilophorus Hahn, and Sthenaridea Reuter (paraphylectic). Pilophorus indonesicus is proposed as a replacement name for Bilirania sumatrana Schuh (=Pilophorus), a secondary homonym of Pilophorus sumatranus Poppius. A list of currently recognized genera and species with summary distributions and host plant associations, and a key to genera are included. Host associations are plotted on the cladogram to reveal the pattern of host shifts. At the generic level and above, a pattern of colonization, rather than co-evolution, is strongly indicated; at the species level, genera of Pilophorini often show restricted plant-group associations, but no clear pattern of coevolution emerges. Major distributional patterns in the Philophorini are mapped, discussed and compared with historical biogeographic schemes for some other groups. The Pilophorini appear to be of tropical Gondwanan origin with subsequent spread into, and differentiation in, the temperate Northern Hemisphere.

HISTORICAL BIOGEOGRAPHY IN THE INDO-PACIFIC: A CLADISTIC APPROACH.

Abstract- The history of biogeographic theories about the Indo-Pacific is reviewed. The methods of cladistic biogeography are introduced and 10 monophyletic groups of Miridae (Insecta: Heteroptera) are used to apply these methods in the Indo-Pacific. The results are summarized and then combined with those of Duffels on cicadas. The results of most previous schemes are rejected on the basis of new results because the prior studies applied methods which failed to correctly interrelate areas of endemism within the Indo-Pacific and between this and other regions. The composite biotic nature of New Guinea is stressed. "it is the presence in and not the absence from land areas of organisms that calls for reasoned explanations" (Evans, 1959).

Integrating specimen databases and revisionary systematics.

Arguments are presented for the merit of integrating specimen databases into the practice of revisionary systematics. Work flows, data connections, data outputs, and data standardization are enumerated as critical aspects of such integration. Background information is provided on the use of "barcodes" as unique specimen identifiers and on methods for efficient data capture. Examples are provided on how to achieve efficient workflows and data standardization, as well as data outputs and data integration.