Phylogeny and age of cockroaches: a reanalysis of mitogenomes with selective fossil calibrations

Abstract

In spite of big data and new techniques, the phylogeny and timing of cockroaches remain in dispute. Apart from sequencing more species, an alternative way to improve the phylogenetic inference and time estimation is to improve the quality of data, calibrations and analytical procedure. This study emphasizes the completeness of data, the reliability of genes (judged via alignment ambiguity and substitution saturation), and the justification for fossil calibrations. Based on published mitochondrial genomes, the Bayesian phylogeny of cockroaches and termites is recovered as: Corydiinae + (((Cryptocercidae + Isoptera) + ((Anaplectidae + Lamproblattidae) + (Tryonicidae + Blattidae))) + (Pseudophyllodromiinae + (Ectobiinae + (Blattellinae + Blaberidae)))). With two fossil calibrations, namely, Valditermes brenanae and Piniblattella yixianensis, this study dates the crown Dictyoptera to early Jurassic, and crown Blattodea to middle Jurassic. Using the ambiguous ‘roachoid’ fossils to calibrate Dictyoptera+sister pushes these times back to Permian and Triassic. This study also shows that appropriate fossil calibrations are rarer than considered in previous studies.

Phylogenetic reconstruction of Corydioidea (Dictyoptera: Blattodea) provides new insights on the placement of Latindiinae and supports the proposal of the new subfamily Ctenoneurinae

“…The genus Arenivaga constitutes a lineage of burrowing cockroaches distributed in North America and Mexico (Hopkins, 2014). In most of our analyses (Dataset 1: ML; Dataset 2: BI and ML), Arenivaga was recovered sister to the rest of the Corydiinae (Figures 1, S2, S4, and S5) with high support values (BS/SH‐aLRT/EQP‐IC = 100/100/94 in Dataset 1; BS/SH‐aLRT/EQP‐IC = 100/100/100, BPP = 1.0), similar to the results of Bourguignon et al (2018) and Li (2022). When analysing Dataset 1 under BI, Arenivaga was recovered sister to other Polyphagini species but with a low support (BPP = 0.51, Figure S3).…”

“…Two of these fossil calibrations († Archeorhinotermes rossi Krishna & Grimaldi and † Valditermes brenanae Jarzembowski) were previously defined and used in Evangelista et al (2019). A third calibration consists of the fossil species † Piniblattella yixianensis Gao, Shih and Ren, which is the oldest known Blattellinae and has been shown to be suitable for the calibration of the most recent common ancestor (MRCA) of Blattellidae and their sister group (Gao et al, 2018; Li, 2022). The fourth calibration consists of the fossil species † Cretaperiplaneta kaonashi Qiu, Che and Wang, which has external morphologic characters typical of Blattidae and shares some features with Tryonicidae, indicating that it can be used as a fossil calibration for the MRCA of Blattidae + Tryonicidae (Deng et al, 2023; Qiu et al, 2020a).…”

“…This species has been previously used as a calibration to set a minimum age for the MRCA of several corydiid nodes (Evangelista et al, 2019; Liu et al, 2023; Wang et al, 2017). However, its taxonomic assignment is doubtful according to Li (2022). To investigate its influence on the age and divergence time of Corydioidea, we performed the dating analyses with or without using † Cretaholocompsa montsecana as a fossil constraint.…”

“…Owing to differences in terms of fossil calibrations and taxon sampling, the estimated age of Corydioidea varies considerably between studies, from late Permian (Djernæs et al, 2015) to early Jurassic (Liu et al, 2023). In addition, Corydioidea were under‐sampled in previous studies, especially the Nocticolidae and Latindiinae (Bourguignon et al, 2018; Djernæs et al, 2015; Evangelista et al, 2019; Legendre et al, 2015; Li, 2022; Li & Huang, 2020; Liu et al, 2023; Wang et al, 2017). Therefore, the analysis of a more comprehensive molecular dataset with relevant fossil calibrations is expected to improve the accuracy of divergence time estimates for Corydioidea (Ksepka et al, 2015; Wolfe et al, 2016).…”

Phylogenetic analysis of Blaberoidea reveals non‐monophyly of taxa and supports the creation of multiple new subfamilies

“…The sister group of the speciose family Blaberidae has remained ambiguous, with a number of candidates proposed, including Pseudophyllodromiidae (Grandcolas, 1996), Ectobiidae + Pseudophyllodromiidae (Djernaes et al, 2020), Blattellinae (Bourguignon et al, 2018;Li, 2022), Nyctiborinae (Klass, 1997;Wang et al, 2017a) and Nyctiboridae + Blattellidae (Evangelista et al, 2019(Evangelista et al, , 2021. At present, there are 12 subfamilies within Blaberidae, including 10 subfamilies recorded by Roth (2003), Paranauphoetinae elevated by Anisyutkin (2003) and Geoscapheinae proposed by Rugg and Rose (1984).…”