Hans Pohl

Morphological and molecular evidence converge upon a robust phylogeny of the megadiverse Holometabola

We present the largest morphological character set ever compiled for Holometabola. This was made possible through an optimized acquisition of data. Based on our analyses and recently published hypotheses based on molecular data, we discuss higher‐level phylogeny and evolutionary changes. We comment on the information content of different character systems and discuss the role of morphology in the age of phylogenomics. Microcomputer tomography in combination with other techniques proved highly efficient for acquiring and documenting morphological data. Detailed anatomical information (356 characters) is now available for 30 representatives of all holometabolan orders. A combination of traditional and novel techniques complemented each other and rapidly provided reliable data. In addition, our approach facilitates documenting the anatomy of model organisms. Our results show little congruence with studies based on rRNA, but confirm most clades retrieved in a recent study based on nuclear genes: Holometabola excluding Hymenoptera, Coleopterida (= Strepsiptera + Coleoptera), Neuropterida excl. Neuroptera, and Mecoptera. Mecopterida (= Antliophora + Amphiesmenoptera) was retrieved only in Bayesian analyses. All orders except Megaloptera are monophyletic. Problems in the analyses are caused by taxa with numerous autapomorphies and/or inapplicable character states due to the loss of major structures (such as wings). Different factors have contributed to the evolutionary success of various holometabolan lineages. It is likely that good flying performance, the ability to occupy different habitats as larvae and adults, parasitism, liquid feeding, and co‐evolution with flowering plants have played important roles. We argue that even in the “age of phylogenomics”, comparative morphology will still play a vital role. In addition, morphology is essential for reconstructing major evolutionary transformations at the phenotypic level, for testing evolutionary scenarios, and for placing fossil taxa.
© The Willi Hennig Society 2010.

Genomic and Morphological Evidence Converge to Resolve the Enigma of Strepsiptera

The phylogeny of insects, one of the most spectacular radiations of life on earth, has received considerable attention. However, the evolutionary roots of one intriguing group of insects, the twisted-wing parasites (Strepsiptera), remain unclear despite centuries of study and debate. Strepsiptera exhibit exceptional larval developmental features, consistent with a predicted step from direct (hemimetabolous) larval development to complete metamorphosis that could have set the stage for the spectacular radiation of metamorphic (holometabolous) insects. Here we report the sequencing of a Strepsiptera genome and show that the analysis of sequence-based genomic data (comprising more than 18 million nucleotides from nearly 4,500 genes obtained from a total of 13 insect genomes), along with genomic metacharacters, clarifies the phylogenetic origin of Strepsiptera and sheds light on the evolution of holometabolous insect development. Our results provide overwhelming support for Strepsiptera as the closest living relatives of beetles (Coleoptera). They demonstrate that the larval developmental features of Strepsiptera, reminiscent of those of hemimetabolous insects, are the result of convergence. Our analyses solve the long-standing enigma of the evolutionary roots of Strepsiptera and reveal that the holometabolous mode of insect development is more malleable than previously thought.

Endopterygote systematics – where do we stand and what is the goal (Hexapoda, Arthropoda)?

Abstract.  Impressive progress has been made recently in the systematics of holometabolous insects. Nevertheless, important questions remain controversial, and uncertainties concerning the relationships of major lineages may even have increased. New analytical techniques have been developed and an immense wealth of molecular data has accumulated. Although no decisive breakthrough has yet been achieved, recent analyses of large molecular datasets have contributed greatly to the reconstruction of the phylogeny of several holometabolous lineages. Extensive combined analyses with substantial morphological datasets and molecular data comprising several genes (‘total evidence’) are still required for a well‐founded phylogenetic hypothesis for the entire group. Endopterygota monophyly is supported mainly by the specific and derived mode of development, which may be considered as a new evolutionary level within Hexapoda. The basal branching pattern remains controversial. A division into two large clades comprising Coleoptera (+ Strepsiptera?) + Neuropterida, on the one hand, and Hymenoptera + [Amphiesmenoptera + Antliophora (including Strepsiptera?)], on the other, appears plausible. Alternative hypotheses have been proposed based on wing characters and molecular data. The position of Strepsiptera remains unsolved. Mecoptera almost certainly is not monophyletic, as Siphonaptera are probably sister to Boreidae. Immense progress has been made in the reconstruction of the intraordinal relationships of all orders, thanks to increasing studies based on combined datasets. Common efforts of morphologists and molecular systematists probably will lead to further rapid progress. Several discoveries of new higher ranking taxa during recent years have revealed that large‐scale habitat destruction will not only have disastrous effects on global biodiversity, but also on the study of insect phylogeny and evolution.

A scanning electron microscopy specimen holder for viewing different angles of a single specimen.

The specimen holder for scanning electron microscopy described herein allows a single specimen to be examined in any possible view and significantly improves object illumination. The specimen is glued to a fine pin and flexibly mounted on a double‐sided adhesive conductive pad on a rotatable pivot. A milled pot placed beneath the specimen acts as an electron trap. This provides a homogeneous black image background by minimizing noisy signals from the specimens surroundings. Microsc. Res. Tech. 73:1073–1076, 2010.

The phylogeny of Strepsiptera (Hexapoda)

Previous phylogenetic analyses of Strepsiptera have been limited to characters from only males or first instar larvae, and by poor taxonomic sampling. This investigation is the first cladistic analysis to use more than fourfold as many characters as any prior study, and a broader sampling of taxa. The analysis of 189 morphological characters of all stages of representatives of all extant strepsipteran families and characters of adult males of amber fossils results in the following branching pattern: (†Protoxenos+ (†Cretostylops + (†Mengea + (Mengenillidae + (Corioxenidae + (Bohartillidae + (Halictophagidae + (Elenchidae + (†Protelencholax + (Myrmecolacidae + (Callipharixenidae + (Xenidae + Stylopidae)))))))))))). The basal placement of the Baltic amber fossil †Protoxenos and the Burmese amber fossil †Cretostylops is well founded. Even though †Cretostylops is older than †Protoxenos it is almost certainly not the most basal strepsipteran group but the sister group of a clade comprising the Baltic amber fossil †Mengea + Strepsiptera s. str. (excl. stemlineage). Monophyly of Mengenillidae, Stylopidia, Stylopiformia s.l., Corioxenidae, Xenidae, and Stylopidae is confirmed. Mengenillidia is paraphyletic (with respect to †Mengea (Mengeidae)), Elenchidae (with respect to †Protelencholax) and the genus Stichotrema (with respect to the Baltic amber fossils). Thus Protelencholacidae fam. n. is described, and S. weitschati and S. triangulum are transferred to Palaeomyrmecolax. A ground plan of adult male Strepsiptera is provided and evolutionary interpretations are presented based on the obtained cladograms.

Insect morphology in the age of phylogenomics: innovative techniques and its future role in systematics

A brief account of the history of insect morphology is given. Different techniques and analytical methods used in current projects on insect morphology and phylogeny and their optimized combined application are described. These include fixation, dissection, maceration, histology (microtome sectioning), scanning electron microscopy (SEM), transmission electron microscopy (TEM), serial block‐face scanning electron microscopy (SBFSEM), focused ion beam scanning electron microscopy (FIB/SEM), confocal laser scanning microscopy (CLSM), bleaching, micro‐computed tomography (μCT), computer‐based three‐dimensional reconstruction, focus stacking of digital images, geometric morphometrics and the storage of morphological metadata. The role of insect morphology in the “age of phylogenomics” is discussed.

Resolving insect phylogeny: The significance of cephalic structures of the Nannomecoptera in understanding endopterygote relationships.

The Nannomecoptera are among the most enigmatic and controversial taxa of endopterygote insects, the phylogenetic resolution of which is crucial to understanding the evolution of neopteran insects. Once considered a subordinate lineage within the Mecoptera, renewed interest in nannochoristids has been prompted by evidence that the Nannomecoptera are not admissible to the clade of extant scorpion flies but are more likely to belong to the clade Siphonaptera + Nannomecoptera + Diptera. The overarching purpose of the present account is to provide novel and extensive morphological character traits in addition to those already existing for adult structures. The aim is to determine if these traits support molecular data sets that have been suggested elsewhere to clarify the phyletic position of Nannochoristidae. This account focuses on nannomecopteran larvae, which unlike those of other mecopterans have received little attention. Thus, the thrust of this investigation is to provide detailed anatomical data on nannochoristid larvae for a targeted inquiry into their phylogenetic affinities. The described characters are discussed and presented in a data matrix comprising representatives of all endopterygote orders. While the study is based primarily on the New Zealand species Nannochorista philpotti, it is proposed that all nannomecopteran larvae will prove to be similar to this taxon in most if not all structural features of significance to a higher-level phylogenetic context.

Protoxenidae fam. nov. (Insecta, Strepsiptera) from Baltic amber — a ‘missing link’ in strepsipteran phylogeny

A male specimen of a new strepsipteran genus and species (Protoxenos janzeni gen. et sp. nov.) and family (Protoxenidae fam. nov.) found in Baltic amber is described and illustrated. It shows features which are apparently more plesiomorphic than in hitherto known strepsipterans, such as laterally inserted eight‐segmented antennae, very robust mandibles with a broad base, a prominent galea, a comparatively short, transverse metapostnotum, hindwings that are feebly extended in a rostrocaudal direction, and equally sclerotized abdominal tergites and sternites. Based on a cladistic analysis of 46 characters of males of 11 genera and three outgroup taxa, P. janzeni is the sister group of all other known strepsipterans, and Mengea the sister group of Strepsiptera s.s. Eoxenos is the sister group of the remaining extant strepsipterans and Mengenillidae is therefore paraphyletic. Newly established groundplan features of Strepsiptera will facilitate the clarification of the systematic position of the Order in future studies.

Phylogeny of the Strepsiptera based on morphological data of the first instar larvae

This investigation was the first cladistic analysis using morphological data of first instar larvae of Strepsiptera. The analysis of representatives of nearly all known families of Strepsiptera supports the division of Strepsiptera into Mengenillidia and Stylopidia. Corioxenidae and Elenchidae are placed at the base of Stylopidia. Halictophagidae is the sister group to Xeninae + Myrmecolacidae + Stylopinae. Xeninae is placed as the sister group to Myrmecolacidae + Stylopinae. Stylopidae are paraphyletic. Thus, Xenidae stat. n. is re‐established. A sister‐group relationship between Myrmecolacidae and Elenchidae is not supported on characters of first instar larvae.

Bahiaxenidae, a “living fossil” and a new family of Strepsiptera (Hexapoda) discovered in Brazil

An adult male of a newly discovered strepsipteran species from Brazil—Bahiaxenos relictus—is described. A new family Bahiaxenidae is suggested based on cladistic analyses of comprehensive morphological data sets with a broad taxon sampling including the stem group. It is unambiguously placed as the sister group of all other extant families of Strepsiptera. Bahiaxenos relictus is the only species of basal, i.e. non‐stylopidian, Strepsiptera occurring in the New World. It appears to be a relict taxon that has survived in the fossil sand dunes of the São Francisco River (Bahia State). The loss of the 8th antennomere and the greatly reduced labrum are autapomorphies of Strepsiptera s.s. excluding Bahiaxenidae. The sister group relationship between†Protoxenidae and the remaining Strepsiptera, and between †Cretostylops and a clade comprising †Mengea and Strepsiptera s.s., is confirmed, as is the monophyly of Stylopidia and Stylopiformia.