Phylogeny of cockroaches (Insecta, Dictyoptera, Blattodea), with placement of aberrant taxa and exploration of out‐group sampling

Abstract

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.

Molecular phylogeny of Blaberidae (Dictyoptera, Blattodea) with implications for taxonomy and evolutionary studies

“…These unsurprising results have been suggested several times and concern the monophyly of Blaberidae, (Geoscapheinae + Panesthiinae) and Oxyhaloinae (Maekawa et al 2003;Inward 2007;Pellens et al 2007;Djernaes et al 2012;Legendre et al 2014Legendre et al , 2015. Likewise, a strong support was found, here and in previous studies, for Gyninae, Panchlorinae and Pycnoscelinae.…”

“…Later, cockroach systematists identified several morphological synapomorphies supporting the monophyly of Blaberidae (Grandcolas 1996;Klass & Meier 2006; brood sac underneath the female genitalia, sclerite R3v small, first vein of vannus with many basal rami, tracheation very dense and tracheae vesicle-shaped, presence of a muscle connecting sclerite L2 and pouch lve). Monophyly has also repeatedly been found with molecular data as well (Maekawa et al 2003;Inward 2007;Pellens et al 2007;Djernaes et al 2012;Legendre et al 2014Legendre et al , 2015.…”

“…1), one Corydiidae (Therea petiveriana Linnaeus, 1758) and one Blattidae (Blatta orientalis Linnaeus, 1758) as the rooting outgroup taxon. Among Ectobiidae, all subfamilies have been postulated as being the sister group of Blaberidae, including Pseudophyllodromiinae (Grandcolas 1996;Inward et al 2007;Pellens et al 2007;Djernaes et al 2012;Legendre et al 2015). At first, we included several species of Pseudophyllodromiinae (Anisopygia sp., Asemoblattana sp., Euphyllodromia sp., Latiblattella sp.…”

The complete mitogenome of the wood-feeding cockroachCryptocercus kyebangensis(Blattodea: Cryptocercidae) and phylogenetic relations among cockroach families

“…Thus, these results strongly support the paraphyly of Blattodea in relation to termites, as suggested in other studies (e.g. Lo et al 2000;Pellens et al 2007;Legendre et al 2008;Cameron et al 2012;Djernaes et al 2012Djernaes et al , 2015. The current phylogenetic tree supports the sister relation of Blattidae and Blattellidae, with the clade Cryptocercidae + Isoptera as a sister group.…”

“…In addition, recent phylogenetic studies involving various mitochondrial or/and nuclear molecular markers showed that Cryptocercus is a sister group of a primitive termite species, Mastotermes darwiniensis (Lo et al 2000;Legendre et al 2008;Cameron et al 2012;Djernaes et al 2012). Because of these behavioral characteristics and their close relation with termites, wood-eating cockroaches of Cryptocercus have often been used as model species to elucidate the early stages of social evolution in termites (Klass et al 2008;Djernaes et al 2015).…”