Revision of the genus Attaphila (Blattodea: Blaberoidea), myrmecophiles living in the mushroom gardens of leaf-cutting ants

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Abstract

The genus Attaphila, comprising minute myrmecophilous cockroaches, is revised, including now six previously known (A. aptera, A. bergi, A. flava, A. fungicola, A. schuppi, A. sexdentis) and three new species (A. multisetosa sp. nov. Bohn and Klass, A. paucisetosa sp. nov. Bohn and Klass, A. sinuosocarinata sp. nov. Bohn and Klass). All species are described or redescribed and depicted with their main characteristics; determination keys allow the identification of males and females. Especially the male characters allow a distribution to two species-groups with differing host specificity: bergi-group associated with Acromyrmex (and possibly Amoimyrmex) ants, fungicola-group associated with Atta ants; the former appears paraphyletic, the latter monophyletic. The genus Attaphila is characterised emphasising its unique features: (1) insertion of antennae at the bottom of a wide funnel-shaped deepening; (2) antenna with the possibility of a rectangular bending between scapus and pedicellus (associated with a distal excavation of the scapus) and (3) with an unusual shape and low number of antennomeres; (4) femora of legs with a ventral groove allowing a close spacing of femur and tibia during a strong flexion; (5) a complex and unusual shape of the laterosternal shelf area of the female genitalia (lack of shelf, presence of a pair of complicated tubular invaginations); and (6) lateral parts of abdominal tergite T9 of male ending in a pair of ventromesally directed arms, which contact the lateral margins of the subgenital plate. Functional aspects and the possible biological roles of these features are discussed. Older biological data are summarised and new observations are presented. The position of Attaphila within Blattodea is discussed. Like a recent molecular study, the morphology of the male genitalia places the genus in the Blaberoidea. The molecular result of Attaphila being closest to three particular blattellid genera, however, is conflictual from the morphological perspective.

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Systematics and phylogeny of cockroaches (Dictyoptera: Blattaria)

Louis M. Roth (2003)

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Phylogeny of Dictyoptera: Dating the Origin of Cockroaches, Praying Mantises and Termites with Molecular Data and Controlled Fossil Evidence

Frédéric Legendre, André Nel, Gavin Svenson … (2015)

Understanding the origin and diversification of organisms requires a good phylogenetic estimate of their age and diversification rates. This estimate can be difficult to obtain when samples are limited and fossil records are disputed, as in Dictyoptera. To choose among competing hypotheses of origin for dictyopteran suborders, we root a phylogenetic analysis (~800 taxa, 10 kbp) within a large selection of outgroups and calibrate datings with fossils attributed to lineages with clear synapomorphies. We find the following topology: (mantises, (other cockroaches, (Cryptocercidae, termites)). Our datings suggest that crown-Dictyoptera—and stem-mantises—would date back to the Late Carboniferous (~ 300 Mya), a result compatible with the oldest putative fossil of stem-dictyoptera. Crown-mantises, however, would be much more recent (~ 200 Mya; Triassic/Jurassic boundary). This pattern (i.e., old origin and more recent diversification) suggests a scenario of replacement in carnivory among polyneopterous insects. The most recent common ancestor of (cockroaches + termites) would date back to the Permian (~275 Mya), which contradicts the hypothesis of a Devonian origin of cockroaches. Stem-termites would date back to the Triassic/Jurassic boundary, which refutes a Triassic origin. We suggest directions in extant and extinct species sampling to sharpen this chronological framework and dictyopteran evolutionary studies.

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An integrative phylogenomic approach illuminates the evolutionary history of cockroaches and termites (Blattodea)

Dominic A. Evangelista, Benjamin Wipfler, Olivier Béthoux … (2019)

Phylogenetic relationships among subgroups of cockroaches and termites are still matters of debate. Their divergence times and major phenotypic transitions during evolution are also not yet settled. We addressed these points by combining the first nuclear phylogenomic study of termites and cockroaches with a thorough approach to divergence time analysis, identification of endosymbionts, and reconstruction of ancestral morphological traits and behaviour. Analyses of the phylogenetic relationships within Blattodea robustly confirm previously uncertain hypotheses such as the sister-group relationship between Blaberoidea and remaining Blattodea, and Lamproblatta being the closest relative to the social and wood-feeding Cryptocercus and termites. Consequently, we propose new names for various clades in Blattodea: Cryptocercus + termites = Tutricablattae; Lamproblattidae + Tutricablattae = Kittrickea; and Blattoidea + Corydioidea = Solumblattodea. Our inferred divergence times contradict previous studies by showing that most subgroups of Blattodea evolved in the Cretaceous, reducing the gap between molecular estimates of divergence times and the fossil record. On a phenotypic level, the blattodean ground-plan is for egg packages to be laid directly in a hole while other forms of oviposition, including ovovivipary and vivipary, arose later. Finally, other changes in egg care strategy may have allowed for the adaptation of nest building and other novelties.