Abstract. Morphological characters and molecular sequence data were for the first time analysed separately and combined for the true water bugs (Hemiptera–Heteroptera, infraorder Nepomorpha). Data from forty species representing all families were included, together with two outgroup species representing the infraorders Gerromorpha and Leptopodomorpha. The morphological data matrix consisted of sixty‐five characters obtained from literature sources. Molecular data included approximately 960 bp from the mitochondrial gene 16S and the nuclear gene 28S for all forty‐two terminal taxa. The morphological dataset was analysed using maximum parsimony and the combined morphological and molecular (16S + 28S rDNA) dataset was analysed using direct optimization. A sensitivity analysis of sixteen different sets of parameters (various combinations of insertion–deletion cost and transversion costs) was undertaken. Character congruence was used as an optimality criterion to choose among competing phylogenetic hypotheses. The final hypothesis was obtained from the analysis of the combined molecular and mor phological dataset with the most congruent parameter set. This hypothesis supports the monophyly of all currently recognized families of Nepomorpha, and of the superfamilies Nepoidea (Nepidae + Belostomatidae), Corixoidea (Corixidae), Ochteroidea Ochteridae + Gelastocoridae), Notonectoidea (Notonectidae), and Pleoidea (Pleidae + Helotrephidae), but not the monophyly of the Naucoroidea (Naucoridae + Aphelocheiridae + Potamocoridae). The close relationship between the Notonectidae and Pleoidea is also supported. Our hypothesis concurs with Mahner in the placement of the Corixidae as a sister group to the remaining nepomorphan superfamilies except the Nepoidea, but differs in the placement of the Ochteroidea as a sister group to the Notonectoidea + Pleoidea. The superfamily Naucoroidea should be limited to only including the family Naucoridae and not the families Aphelocheiridae and Potamocoridae. The present analysis strongly supports a sister group relationship between the families Aphelocheiridae and Potamocoridae, a monophylum for which we propose a new superfamily, Aphelocheiroidea.
The phylogeny of semi-aquatic bugs (Hemiptera-Heteroptera: Gerromorpha) was tested in parsimony analyses of 64 morphological characters and approximately 2.5 kb of DNA sequence data from the mitochondrial genes encoding COI+II and 16SrRNA and the nuclear gene encoding 28SrRNA. The taxon sample included representatives of all families and most subfamilies of Gerromorpha and a selection of outgroup taxa representing the two basal infraorders of Heteroptera, Enicocephalomorpha and Dipsocoromorpha, and two families of Nepomorpha. A simultaneous analysis (SA) of all data, and with gaps scored as fifth state characters, gave a single most parsimonious tree with all families resolved as monophyletic, except the Veliidae, where Microveliinae + Haloveliinae, Veliinae, Rhagoveliinae, Perittopinae, and Ocelloveliinae were resolved as successive sister groups to the Gerridae, thus confirming earlier statements about paraphyly of this family. The Gerridae + Veliidae clade was strongly supported, but otherwise only the Gerridae + Veliidae less Ocelloveliinae and the Gerridae itself had support. These three clades could all be diagnosed on apomorphic morphological characters, although no characters diagnosing the Gerridae were without convergences or present in all included taxa. While the Ocelloveliinae, Veliinae and Haloveliinae could not be diagnosed on convincing apomorphies, the Microveliinae + Haloveliinae, and their sister group relationship with the Gerridae, could be diagnosed on rather strong morphological synapomorphies, suggesting that Gerridae could be expanded to include these two veliid subfamilies, while Ocelloveliinae, and perhaps the remaining veliid subfamilies, could be elevated to new families. In Gerridae, the Ptilomerinae + Halobatinae was sister group to all other subfamilies, while the Rhagadotarsinae + Trepobatinae was sister group to a clade comprising the Gerrinae, Eotrechinae, Cylindrostethinae and Charmatometrinae. Most relationships in this clade were poorly supported and diagnosed, and Cylindrostethinae was surprisingly found to be paraphyletic. The sister group to the Gerridae + Veliidae clade was a strongly supported clade comprising the Paraphrynoveliidae and Macroveliidae, and this, and the lack of convincing synapomorphies for Paraphrynoveliidae, suggest that these two small families could be synonymized. For the basal relationships of Gerromorpha, the Mesoveliidae was strongly supported sister group to all other families, while the Hebridae, Hermatobatidae and Hydrometridae formed a poorly supported and poorly diagnosed sister group to the Gerridae + Veliidae + Paraphrynoveliidae + Macroveliidae clade. The unexpected sister group relationship between Hermatobatidae and Hydrometridae was moderately supported, and could be diagnosed on two synapomorphies, thus giving a new hypothesis about the relationships of these very divergent families. Phylogenetic analyses of individual character partitions gave less resolved and less supported relationships, and the mitochondrial genes COI+II and 16SrRNA contributed negative hidden partitioned Bremer support (HPBS) to the simultaneous analysis tree, probably due to homoplasy caused by saturation effects.
Heteroptera are among the most diverse hemimetabolous insects. Seven infraorders have been recognized within this suborder of Hemiptera. Apart from the well-established sister-group relationship between Cimicomorpha and Pentatomomorpha (= Terheteroptera), the two terminal lineages, the relationships among the other five infraorders are still controversial, of which three (Gerromorpha, Nepomorpha and Leptopodomorpha) are intimately connected to aquatic environments. However, the various and often conflicting available phylogeny hypotheses do not offer a clear background for a connection between diversification and palaeoenvironments. In this study, a molecular data set representing 79 taxa and 10 149 homologous sites is used to infer the phylogenetic relationships within Heteroptera. Bayesian inference, maximum-likelihood and maximum parsimony analyses were employed. The results of phylogenetic inferences largely confirm the widely accepted phylogenetic context. Estimation of the divergence time based on the phylogenetic results revealed that Gerromorpha, Nepomorpha and Leptopodomorpha originated successively during the period from the Late Permian to Early Triassic . This timescale is consistent with the origin and radiation time of various aquatic holometabolans. Our results indicate that the aquatic and semi-aquatic true bugs evolved under environmental conditions of high air temperature and humidity in an evolutionary scenario similar to that of the aquatic holometabolans.
We addressed the phylogeny of cockroaches using DNA sequence data from a broad taxon sample of Dictyoptera and other non‐endopterygotan insect orders. We paid special attention to several taxa in which relationships are controversial, or where no molecular evidence has been used previously: Nocticolidae, a family of small, often cave‐dwelling cockroaches, has been suggested to be the sister group of the predaceous Mantodea or of the cockroach family Polyphagidae; Lamproblatta, traditionally placed in Blattidae, has recently been given family status and placed as sister to Polyphagidae; and Saltoblattella montistabularis Bohn, Picker, Klass & Colville, a jumping cockroach, which has not yet been included in any phylogenetic studies. We used mitochondrial (COI + COII and 16S) and nuclear (18S and 28S) genes, and analysed the data using Bayesian inference (BI) and maximum likelihood (ML). Nocticolidae was recovered as sister to Polyphagidae. Lamproblatta was recovered as sister to Blattidae, consistent with the traditional placement (not based on phylogenetic analysis). However, because of the limited support for this relationship and conflict with earlier morphology‐based phylogenetic hypotheses, we retain Lamproblattidae. S. montistabularis was consistently placed as sister to Ectobius sylvestris Poda (Blaberoidea: Ectobinae), indicating that the saltatorial hindlegs of this genus are a relatively recent adaptation. Isoptera was placed within Blattodea as sister to Cryptocercidae. Nocticolidae + Polyphagidae was sister to Isoptera + Cryptocercidae, and Blaberoidea was sister to the remaining Blattodea.
The phylogeny of true bugs (Hemiptera: Heteroptera), one of the most diverse insect groups in terms of morphology and ecology, has been the focus of attention for decades with respect to several deep nodes between the suborders of Hemiptera and the infraorders of Heteroptera. Here, we assembled a phylogenomic data set of 53 taxa and 3102 orthologous genes to investigate the phylogeny of Hemiptera-Heteroptera, and both concatenation and coalescent methods were used. A binode-control approach for data filtering was introduced to reduce the incongruence between different genes, which can improve the performance of phylogenetic reconstruction. Both hypotheses (Coleorrhyncha + Heteroptera) and (Coleorrhyncha + Auchenorrhyncha) received support from various analyses, in which the former is more consistent with the morphological evidence. Based on a divergence time estimation performed on genes with a strong phylogenetic signal, the origin of true bugs was dated to 290-268 Ma in the Permian, the time in Earth's history with the highest concentration of atmospheric oxygen. During this time interval, at least 1007 apomorphic amino acids were retained in the common ancestor of the extant true bugs. These molecular apomorphies are located in 553 orthologous genes, which suggests the common ancestor of the extant true bugs may have experienced large-scale evolution at the genome level.
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