Cyrille A. D'Haese

Mitochondrial genome data alone are not enough to unambiguously resolve the relationships of Entognatha, Insecta and Crustacea sensu lato (Arthropoda)

An analysis of the relationships of the major arthropod groups was undertaken using mitochondrial genome data to examine the hypotheses that Hexapoda is polyphyletic and that Collembola is more closely related to branchiopod crustaceans than insects. We sought to examine the sensitivity of this relationship to outgroup choice, data treatment, gene choice and optimality criteria used in the phylogenetic analysis of mitochondrial genome data. Additionally we sequenced the mitochondrial genome of an archaeognathan, Nesomachilis australica, to improve taxon selection in the apterygote insects, a group poorly represented in previous mitochondrial phylogenies. The sister group of the Collembola was rarely resolved in our analyses with a significant level of support. The use of different outgroups (myriapods, nematodes, or annelids + mollusks) resulted in many different placements of Collembola. The way in which the dataset was coded for analysis (DNA, DNA with the exclusion of third codon position and as amino acids) also had marked affects on tree topology. We found that nodal support was spread evenly throughout the 13 mitochondrial genes and the exclusion of genes resulted in significantly less resolution in the inferred trees. Optimality criteria had a much lesser effect on topology than the preceding factors; parsimony and Bayesian trees for a given data set and treatment were quite similar. We therefore conclude that the relationships of the extant arthropod groups as inferred by mitochondrial genomes are highly vulnerable to outgroup choice, data treatment and gene choice, and no consistent alternative hypothesis of Collembolas relationships is supported. Pending the resolution of these identified problems with the application of mitogenomic data to basal arthropod relationships, it is difficult to justify the rejection of hexapod monophyly, which is well supported on morphological grounds.

Evolution on a shaky piece of Gondwana: is local endemism recent in New Caledonia?

New Caledonia is well known as a hot spot of biodiversity whose origin as a land mass can be traced back to the Gondwanan supercontinent. The local flora and fauna, in addition to being remarkably rich and endemic, comprise many supposedly relictual groups. Does the New Caledonian biota date back to Gondwanan times, building up its richness and endemism over 100 Myr or does it result from recent diversifications after Tertiary geological catastrophic events? Here we use a molecular phylogenetic approach to answer this question with the study of the Neocaledonian cockroach genus Angustonicus belonging to the subfamily Tryonicinae from Australia and New Caledonia. Both geological and molecular dating show that the diversification of this group is less than two million years old, whatever the date of its origin itself. This dating is not consistent with hypotheses of Gondwanan richness and endemism in New Caledonian biota. In other terms, local richness and endemism at the specific level are not necessarily related to an old Gondwanan origin of the Neocaledonian groups.

A complete insect from the Late Devonian period

After terrestrialization, the diversification of arthropods and vertebrates is thought to have occurred in two distinct phases, the first between the Silurian and the Frasnian stages (Late Devonian period) (425–385 million years (Myr) ago), and the second characterized by the emergence of numerous new major taxa, during the Late Carboniferous period (after 345 Myr ago). These two diversification periods bracket the depauperate vertebrate Romer’s gap (360–345 Myr ago) and arthropod gap (385–325 Myr ago), which could be due to preservational artefact. Although a recent molecular dating has given an age of 390 Myr for the Holometabola, the record of hexapods during the Early–Middle Devonian (411.5–391 Myr ago, Pragian to Givetian stages) is exceptionally sparse and based on fragmentary remains, which hinders the timing of this diversification. Indeed, although Devonian Archaeognatha are problematic, the Pragian of Scotland has given some Collembola and the incomplete insect Rhyniognatha, with its diagnostic dicondylic, metapterygotan mandibles. The oldest, definitively winged insects are from the Serpukhovian stage (latest Early Carboniferous period). Here we report the first complete Late Devonian insect, which was probably a terrestrial species. Its ‘orthopteroid’ mandibles are of an omnivorous type, clearly not modified for a solely carnivorous diet. This discovery narrows the 45-Myr gap in the fossil record of Hexapoda, and demonstrates further a first Devonian phase of diversification for the Hexapoda, as in vertebrates, and suggests that the Pterygota diversified before and during Romer’s gap.

What does Cryptocercus kyebangensis, n.sp. (Dictyoptera: Blattaria: Polyphagidae) from Korea reveal about Cryptocercus evolution? A study in morphology, molecular phylogeny, and chemistry of tergal glands

Abstract The description of a new species of the woodroach Cryptocercus kyebangensis Grandcolas from South Korea offers the opportunity to bring comparative information within the genus. This species, though morphologically very similar to other East Asian and North American species, presents conspicuous differentiation of both ribosomal genes (sequenced fragments of 12S and 16S) and chemical blends from tergal glands (proportions of linalyl acetate and the alcohol 4, 6, 8-trimethyl-7, 9-undecadien-5-ol, compounds previously identified in females originating from North America). A phylogenetic reconstruction involving Blatta orientalis as an outgroup, Therea petiveriana as a polyphagid relative, C. kyebangensis and 17 North American Cryptocercus populations showed that C. kyebangensis stands as a sister-group of North American Cryptocercus, thus suggesting that one beringian vicariance has taken place in the early differentiation of the genus. Using sequence divergences within the framework of the molecular clock leads to comparable datation estimates for both ribosomal genes, ranging between Miocene and Pliocene for the whole history of the genus Cryptocercus, therefore congruent with previous phylogenetic biogeographical estimates. This new set of data for this East Asian species supports heterobathmy in the genus Cryptocercus and argues against the misleading “primitive taxon” concept. Species nova: Cryptocercus kyebangensis Grandcolas.

An ancient Antarctic endemic genus restored: morphological and molecular support for Gomphiocephalus hodgsoni (Collembola: Hypogastruridae)

Gomphiocephalus hodgsoni Carpenter was only the second collembolon † to be described from the Antarctic continent. It was collected first in 1902 from Granite Harbour, southern Victoria Land, Eastern Antarctica, by the British National Antarctic Expedition (1901–1904). Since then several studies have investigated the distribution, ecology, ecophysiology and molecular composition of the species. Despite two morphological redescriptions and an absence of detailed evolutionary phylogenetic studies, the genus Gomphiocephalus was recently reduced to a subgenus of Schoettella Schäffer. Here, we redescribe the species in detail and use morphological and molecular (cytochrome c oxidase subunit I and 28S) data to indicate its generic relationships within Hypogastruridae. Characters of Gomphiocephalus do not conform with those of any extant genus in the family, including Schoettella. In addition, the only Schoettella species described from the southern hemisphere, Schoettella subcorta Salmon, is shown here to belong in the genus Xenylla. Furthermore, molecular data indicates the genus has no close relationship to any other in Poduromorpha, and in particular Hypogastruridae. Therefore, we restore Gomphiocephalus to generic status. Our results reinforce the already recognized high level of endemism in the Antarctic fauna at both species and generic levels, and emphasise the necessity of using both morphological and molecular data in determining the systematics and evolutionary relationships of the fauna.

The phylogenetic position of early hexapod lineages: morphological data contradict molecular data

Abstract.  A review of different studies on the phylogenetic relationships of the early Hexapoda lineages shows that analyses based on molecular sequence data have led to labile and sometimes incongruous results, introducing doubt as to the reliability of the cladograms as a whole. In a recent analysis using molecular data, the Collembola, usually considered as early branching hexapods, appear to occupy a position outside the assemblage of Crustacea and Insecta, leading to the rejection of the traditional view of hexapod monophyly. However, many morphological features, as well as the results of cladistic analyses based on morphological and developmental information, contradict these conclusions. More generally, it appears that in the present state of the analytical strategies, hypotheses concerning arthropod phylogenies obtained from morphological and developmental criteria and combined analyses involving molecular and morphological data provide more reliable results than those generated by molecular information alone.

Antarctic Tardigrada: a first step in understanding molecular operational taxonomic units (MOTUs) and biogeography of cryptic meiofauna

Recent studies have suggested that some resident Antarctic biota are of ancient origin and may have been isolated for millions of years. The phylum Tardigrada, which is part of the Antarctic terrestrial meiofauna, is of particular interest due to an impressive array of biochemical abilities to withstand harsh environmental conditions. Tardigrades are one of the few widespread Antarctic terrestrial animals that have the potential to be used as a model for evolution and biogeography on the Antarctic continent. We isolated 126 individual tardigrades from four geographically isolated soil samples from two remote nunataks in the Sør Rondane Mountains, Dronning Maud Land, Antarctica. We examined genetic variation among individuals utilising three gene regions: cytochrome c oxidase subunit I gene (COI), 18S rDNA (18S), and the wingless (Wg) gene. Comparison of sequences from worldwide and Antarctic tardigrades indicated long-term survival and isolation over glacially dominated periods in ice-free habitats in the Sør Rondane Mountains.

Mapping characters on a tree with or without the outgroups

A character of special interest in evolutionary studies is usually optimized on a phylogenetic tree, with or without the outgroups employed in that analysis. Both practices are never justified and look like arbitrary choices. Focusing on one example, we draw the conclusion that authors retain or remove outgroups depending on the way these outgroups sample the diversity of states of the character(s) of special interest. The topology without outgroups is often used by authors when different outgroup taxa non‐exhaustively sample the different states of the character of interest outside of the ingroup. This can make the analysis incoherent, because its different steps are not based on the same data matrix (outgroups are removed in the last step). It can provide several incoherent and possibly different patterns for a same character of interest, one issuing from the first step of phylogeny construction and the other resulting from the a posteriori optimization on the truncated topology. Phylogenetic analyses should be designed to minimize this problem, selecting outgroup and ingroup taxa whose diversity of character states is needed for reconstructing the evolutionary history of the character of interest.

Garrouste et al. reply

replying to T. Hörnschemeyer et al. Nature 494, 10.1038/nature11887 (2013)Since the nineteenth century, Devonian insects have repeatedly proven to be something else, the sole exception being Rhyniognatha. Recently the Devonian insect Strudiella devonica has been denied by Hörnschemeyer et al., who could not “confirm the presence of a mandible or of mandibular teeth”.

Phylogeny of the lacebugs (Insecta: Heteroptera: Tingidae) using morphological and molecular data

A phylogeny of the Tingidae is provided on the basis of parsimony analysis with pre‐alignment as well as with direct optimization of 2018 bp from five loci (COI, Leu‐tRNA, COII, 16S and 28S) and a set of 30 morphological characters. The results obtained with either direct optimization or pre‐alignment methods of parsimony clearly show that Cantacaderinae are sister group to (Phatnominae + Tinginae). Several apomorphies, retrieved in different studies, characterize Cantacaderinae, Phatnominae and Tinginae, but only one characterizes the grouping of Phatnominae + Tinginae. In addition, no clear tribal division as currently used was highlighted within the Tinginae. The results are discussed in the light of earlier morphologically based hypotheses.