A molecular phylogeny of Phasmatodea with emphasis on Necrosciinae, the most species‐rich subfamily of stick insects

Abstract

The phasmatodeans or stick and leaf insects are considered to be a mesodiverse insect order with more than 3000 species reported mainly from the tropics. The stick insect subfamily Necrosciinae comprises approximately 700 described species in more than 60 genera from the Oriental and Australian region, forming the most species‐rich subfamily traditionally recognized within Phasmatodea. However, the monophyly of this taxon has never been thoroughly tested and the evolutionary relationships among its members are unknown. We analyse three nuclear (18S and 28S rDNA, histone 3) and three mitochondrial (CO II, 12S and 16S rDNA) genes to infer the phylogeny of 60 species of stick insects that represent all recognized families and major subfamilies sensu Günther and the remarkable diversity within Necrosciinae. Maximum parsimony, maximum likelihood and Bayesian techniques largely recover the same substantial clades, albeit with highly discordant relationships between them. Most members of the subfamily Necrosciinae form a clade. However, the genus Neohirasea – currently classified within Lonchodinae – is strongly supported as subordinate to Necrosciinae, whereas Baculofractum, currently classified within Necrosciinae, is strongly supported within Lonchodinae. Accordingly, we formally transfer Neohirasea and allied taxa (namely Neohiraseini) to Necrosciinae sensu nova (s.n.) and Baculofractum to Lonchodinae s.n. We also provide further evidence that Leprocaulinus, until recently recognized as Necrosciinae, belongs to Lonchodinae, and forms the sister taxon of Baculofractum. Furthermore, Lonchodinae is paraphyletic under exclusion of Eurycantha and Neopromachus. We reinstate the traditional view that Neopromachus and related taxa (Neopromachini sensu Günther) are a subgroup of Lonchodinae and transfer those taxa + the New Guinean Eurycanthinae accordingly. Morphological evidence largely corroborates our molecular‐based findings and also reveals that Menexenus fruhstorferi is a member of the genus Neohirasea and is thus transferred from Menexenus (Lonchodinae) to Neohirasea, as Neohirasea fruhstorferi comb.n. (Necrosciinae s.n.). Other phylogenetic results include Areolatae and Anareolatae each supported as polyphyletic, Heteropteryginae and Lanceocercata (Bayesian analysis) are monophyletic, albeit with low support, and Necrosciinae s.n. and Lonchodinae s.n. are recovered as sister taxa (Bayesian analysis).

Review of the Gross Anatomy and Microbiology of the Phasmatodea Digestive Tract

“…the relatively short and stout timematidae are considered the sister group to the euphasmatodea (Whiting et al 2003, terry & Whiting 2005, Friedemann et al 2012, Gottardo et al 2012, tilgner et al 1999. Recent works suggest phasmatodea are a monophyletic group based on morphological (tilgner et al 1999, Friedemann et al 2012, Beutel et al 2013), mtdnA (plazzi et al 2011, Bradler et al 2014, nuclear genomic (Buckley et al 2009, Bradler et al 2014, and transcriptomic (Misof et al 2014) datasets. Autapomorphies include prothoracic defense glands and small, pyriform appendages on the posterior midgut (tilgner et al 1999(tilgner et al , Beutel et al 2013 recently demonstrated as having a unique excretory function (Shelomi & Kimsey 2014).…”

“…Generality of phasmid gut morphology.-of the eleven species available, anatomical studies were performed on nine, all but two (R. nematodes and M. extradentata) from different subfamilies sensu Bradler et al (2014) (Fig. 1).…”

Morphology of the terminalia of the stick insect Dajaca napolovi from Vietnam (Insecta: Phasmatodea)

“…; Bradler et al. ). The postabdomen of Phasmatodea exhibits various exoskeletal specializations that surround the external genitalia (Bradler , ; Helm et al.…”

“…Phasmatodea (stick and leaf insects) are a mesodiverse order of polyneopteran insects comprising about 3000 described species (Terry & Whiting 2005;Bradler 2015). They consist of two majors extant lineages, Timema and Euphasmatodea (including all remaining taxa) (Whiting et al 2003;Klug & Bradler 2006;Friedemann et al 2012;Bradler et al 2014). The postabdomen of Phasmatodea exhibits various exoskeletal specializations that surround the external genitalia (Bradler , 2009Helm et al 2011).…”

Progress, pitfalls and parallel universes: a history of insect phylogenetics

“…Given their almost unimaginable diversity, it is impossible for any individual to be considered an expert for all Hexapoda, and most workers spend their careers exploring particular groups. Here we list a sample of the recent advances from various authors in the phylogeny of Odonata [283 -290,292], Ephemeroptera [278,293,294], Plecoptera [295], Dermaptera [296], Embioptera [297], Phasmatodea [298], Dictyoptera [236,279,[299][300][301], Mantodea [302], Orthoptera [234,303 -306], Hemiptera [307,308], Psocodea [309], Hymenoptera [310 -317], Neuropterida [318 -320], Coleoptera [321 -323], Diptera [324,325], Lepidoptera [326 -334], Trichoptera [335 -337], Mecoptera [246,338,339] and Siphonaptera [340,341].…”