Phylogeny of the Dictyoptera

Molecular phylogenetics …, 2008

A phylogenetic hypothesis of termite relationships was inferred from DNA sequence data. Seven gene fragments (12S rDNA, 16S rDNA, 18S rDNA, 28S rDNA, cytochrome oxidase I, cytochrome oxidase II and cytochrome b) were sequenced for 40 termite exemplars, representing all termite families and 14 outgroups. Termites were found to be monophyletic with Mastotermes darwiniensis (Mastotermitidae) as sister group to the remainder of the termites. In this remainder, the family Kalotermitidae was sister group to other families. The families Kalotermitidae, Hodotermitidae and Termitidae were retrieved as monophyletic whereas the Termopsidae and Rhinotermitidae appeared paraphyletic. All of these results were very stable and supported with high bootstrap and Bremer values. The evolution of worker caste and foraging behavior were discussed according to the phylogenetic hypothesis. Our analyses suggested that both true workers and pseudergates (‘‘false workers”) were the result of at least two different origins. Our data support a traditional hypothesis of foraging behavior, in which the evolutionary transition from a one-piece type to a separate life type occurred through an intermediate behavioral form.

Entomologische …, 2003

This work presents a critical evaluation of the currently available morphological and biological data base relevant to reconstructing phylogeny in Dictyoptera. In 1992, B.L. Thorne and J.M. Carpenter (Systematic Entomology 17: 253-268) published a widely acknowledged phylogenetic analysis of Dictyoptera that gave the relationships (Mantodea + (*Blattaria + Cryptocercidae)) + ((Mastotermitidae + Kalotermitidae) + Termopsidae) [*Blattaria = Blattaria excluding Cryptocercidae]. The present study uses that work as a starting point to demonstrate the importance of a thorough treatment of characters in phylogenetic reconstruction. Repeating the analysis of Thorne and Carpenter without disputable polarity assumptions produced Mantodea + ((*Blattaria + Cryptocercidae) + ((Mastotermitidae + Kalotermitidae) + Termopsidae)). Analyses after a careful revision of the characters as well as analyses including seven additional characters produced Mantodea + (*Blattaria + (Cryptocercidae + (Mastotermitidae + (Kalotermitidae + Termopsidae)))). The latter result, indicating paraphyly of Blattaria with respect to Isoptera, is in strong contrast to the findings of Thorne and Carpenter, but concurs with some recent morphological and molecular studies. It includes a single acquisition of both symbiotic fat body Blattabacterium and hindgut flagellates within Dictyoptera, and the homology of sociality in Cryptocercidae and Isoptera.

PloS one, 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 po...

Molecular Phylogenetics and Evolution, 2008

A phylogenetic hypothesis of termite relationships was inferred from DNA sequence data. Seven gene fragments (12S rDNA, 16S rDNA, 18S rDNA, 28S rDNA, cytochrome oxidase I, cytochrome oxidase II and cytochrome b) were sequenced for 40 termite exemplars, representing all termite families and 14 outgroups. Termites were found to be monophyletic with Mastotermes darwiniensis (Mastotermitidae) as sister group to the remainder of the termites. In this remainder, the family Kalotermitidae was sister group to other families. The families Kalotermitidae, Hodotermitidae and Termitidae were retrieved as monophyletic whereas the Termopsidae and Rhinotermitidae appeared paraphyletic. All of these results were very stable and supported with high bootstrap and Bremer values. The evolution of worker caste and foraging behavior were discussed according to the phylogenetic hypothesis. Our analyses suggested that both true workers and pseudergates (''false workers") were the result of at least two different origins. Our data support a traditional hypothesis of foraging behavior, in which the evolutionary transition from a one-piece type to a separate life type occurred through an intermediate behavioral form.

Molecular phylogenetics and evolution, 2008

Isoptera are highly specialized cockroaches and are one of the few eusocial insect lineages. Cryptocercus cockroaches have appeared to many as ideal models for inference on the early evolution of termites, due to their possible phylogenetic relationship and several shared key attributes in life history. Recently, Pellens, Grandcolas, and colleagues have proposed the blaberid cockroach Parasphaeria boleiriana to be an alternative model for the early evolution in termites. We compare the usefulness of Cryptocercus and P. boleiriana as models for termite evolution. Cryptocercus and lower Isoptera (1) can both feed on comparatively recalcitrant wood, (2) have an obligate, rich and unique hypermastigid and oxymonadid fauna in the hindgut, (3) transfer these flagellates to the next generation by anal trophallaxis, (4) have social systems that involve long-lasting biparental care, and, finally, (5) are strongly suggested to be sister groups, so that the key attributes (1)-(4) appear to be homologous between the two taxa. On the other hand, P. boleiriana (1) feeds on soft, ephemeral wood sources, (2) shows no trace of the oxymonadid and hypermastigid hindgut fauna unique to Cryptocercus and lower Isoptera, nor does it have any other demonstrated obligate relationship with hindgut flagellates, (3) is likely to lack anal trophallaxis, (4) has only a short period of uniparental brood care, and (5) is phylogenetically remote from the Cryptocercus + Isoptera clade. These facts would argue against any reasonable usage of P. boleiriana as a model for the early evolution of Isoptera or even of the clade Cryptocercus + Isoptera. Cryptocercus thus remains an appropriate model-taxon-by-homology for early termite evolution. As compared to P. boleiriana, some other Blaberidae (such as the Panesthiinae Salganea) appear more useful as model-taxa-by-homoplasy for the early evolution of the Cryptocercus + Isoptera clade, as their brooding behavior is more elaborate than in P. boleiriana.

A phylogenetic hypothesis of termite relationships was inferred from DNA sequence data. Seven gene fragments (12S rDNA, 16S rDNA, 18S rDNA, 28S rDNA, cytochrome oxidase I, cytochrome oxidase II and cytochrome b) were sequenced for 40 termite exemplars, representing all termite families and 14 outgroups. Termites were found to be monophyletic with Mastotermes darwiniensis (Mastotermitidae) as sister group to the remainder of the termites. In this remainder, the family Kalotermitidae was sister group to other families. The families Kalotermitidae, Hodotermitidae and Termitidae were retrieved as monophyletic whereas the Termopsidae and Rhinotermitidae appeared paraphyletic. All of these results were very stable and supported with high bootstrap and Bremer values. The evolution of worker caste and foraging behavior were discussed according to the phylogenetic hypothesis. Our analyses suggested that both true workers and pseudergates (''false workers") were the result of at least two different origins. Our data support a traditional hypothesis of foraging behavior, in which the evolutionary transition from a one-piece type to a separate life type occurred through an intermediate behavioral form.

Molecular Phylogenetics and Evolution, 2008

Isoptera are highly specialized cockroaches and are one of the few eusocial insect lineages. Cryptocercus cockroaches have appeared to many as ideal models for inference on the early evolution of termites, due to their possible phylogenetic relationship and several shared key attributes in life history. Recently, Pellens, Grandcolas, and colleagues have proposed the blaberid cockroach Parasphaeria boleiriana to be an alternative model for the early evolution in termites. We compare the usefulness of Cryptocercus and P. boleiriana as models for termite evolution. Cryptocercus and lower Isoptera (1) can both feed on comparatively recalcitrant wood, (2) have an obligate, rich and unique hypermastigid and oxymonadid fauna in the hindgut, (3) transfer these flagellates to the next generation by anal trophallaxis, (4) have social systems that involve long-lasting biparental care, and, finally, (5) are strongly suggested to be sister groups, so that the key attributes (1)-(4) appear to be homologous between the two taxa. On the other hand, P. boleiriana (1) feeds on soft, ephemeral wood sources, (2) shows no trace of the oxymonadid and hypermastigid hindgut fauna unique to Cryptocercus and lower Isoptera, nor does it have any other demonstrated obligate relationship with hindgut flagellates, (3) is likely to lack anal trophallaxis, (4) has only a short period of uniparental brood care, and (5) is phylogenetically remote from the Cryptocercus + Isoptera clade. These facts would argue against any reasonable usage of P. boleiriana as a model for the early evolution of Isoptera or even of the clade Cryptocercus + Isoptera. Cryptocercus thus remains an appropriate model-taxon-by-homology for early termite evolution. As compared to P. boleiriana, some other Blaberidae (such as the Panesthiinae Salganea) appear more useful as model-taxa-by-homoplasy for the early evolution of the Cryptocercus + Isoptera clade, as their brooding behavior is more elaborate than in P. boleiriana.

Arthropod Structure & Development, 2010

Among insects, eusocial behavior occurs in termites, ants, some bees and wasps. Isoptera and Hymenoptera convergently share social behavior, and for both taxa its evolution remains poorly understood. While dating analyses provide researchers with the opportunity to date the origin of eusociality, fossil calibration methodology may mislead subsequent ecological interpretations. Using a comprehensive termite dataset, we explored the effect of fossil placement and calibration methodology. A combined molecular and morphological dataset for 42 extant termite lineages was used, and a second dataset including these 42 taxa, plus an additional 39 fossil lineages for which we had only morphological data. MrBayes doublet-model analyses recovered similar topologies, with one minor exception (Stolotermitidae is sister to the Hodotermitidae, s.s., in the 42-taxon analysis but is in a polytomy with Hodotermitidae and (Kalotermitidae + Neoisoptera) in the 81-taxon analysis). Analyses using the r8s program on these topologies were run with either minimum/maximum constraints (analysis a = 42-taxon and analysis c = 81-taxon analyses) or with the fossil taxon ages fixed (ages fixed to be the geological age of the deposit from which they came, analysis b = 81-taxon analysis). Confidence intervals were determined for the resulting ultrametric trees, and for most major clades there was significant overlap between dates recovered for analyses A and C (with exceptions, such as the nodes Neoisoptera, and Euisoptera). With the exception of isopteran and eusiopteran node ages, however, none of the major clade ages overlapped when analysis B is compared with either analysis A or C. Future studies on Dictyoptera should note that the age of Kalotermitidae was underestimated in absence of kalotermitid fossils with fixed ages.

Systematic Entomology, 2020

Although dissimilar in their overall appearance and life habits, the praying mantises (Mantodea) and cockroaches (Blattodea, including their eusocial relatives, the termites [Isoptera]) are grouped within the clade Dictyoptera, based on-among other significant characteristics-the laying of eggs in a compound structure called an ootheca. The origin of the Dictyoptera and the currently recognized taxa within is, however, a controversial topic among entomologists. This has resulted from disparities in the divergence age estimates obtained from phylogenetic analyses based on molecular data together with the limited and controversial fossil evidence attributable to these groups. Here, we report two new oothecae ichnospecies found in a Carnian (237 to 227 mya. lowermost Upper Triassic) deposit from Argentina. Morphological comparisons and Scanning Electron Microscope and X-ray Energy Dispersive Spectroscopy analyses of fossil and extant oothecae of mantises and cockroaches were performed in an attempt to solve their systematic placement within Dictyoptera and fossil allies, such as †Alienoptera. In addition to being the earliest known record of oothecae, this discovery moves the origin of this specialized reproductive strategy back by 100 million years. As direct fossil evidence, these specimens provide an important calibration and reference point that can inform future research on the origins and timing of diversification of the Dictyoptera.

Biology Letters, 2007