Abstract
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.
Highlights
The hemipteran suborder Heteroptera are part of the most diversified group of non-endopterygote and nonholometabolous insects, including more than 40,000 described species [1].Classification of the Heteroptera, or true bugs, has reached its present state through a long evolutionary process beginning, insofar as modern systematics is concerned, with the work of Linnaeus
RDNA data partition of 1982 characters, a 28S rDNA data partition of 607 characters, a 16S rDNA data partition of 530 characters, a Cytochrome Oxidase Subunit I (COI) data partition of 1067 characters; a nuclear data partition (18S rDNA+28S rDNA) of 2589 characters; a mitochondria data partition (16S rDNA+COI) of 1597 characters; and a total combined data set of 4186 characters
The parsimony results with gaps scored as a fifth character state for both the nuclear data and four-gene combined data were in File S1
Summary
The hemipteran suborder Heteroptera are part of the most diversified group of non-endopterygote and nonholometabolous insects, including more than 40,000 described species [1].Classification of the Heteroptera, or true bugs, has reached its present state through a long evolutionary process beginning, insofar as modern systematics is concerned, with the work of Linnaeus. In the system of Linnaeus, the true bugs were placed in the Hemiptera, the first recognized higher group in Insecta, to include Thysanoptera (thrips), and the other hemipteran suborders Sternorrhyncha (aphids, coccoids) and Auchenorrhyncha (cicadas). Leston et al (1954) [6] introduced the terms Cimicomorpha and Pentatomomorpha in the first formal attempt to recognize natural groups within the Geocorisae.
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