Benjamin Wipfler

• Dr.
• Research Museum Alexander Koenig

In the arid landscapes of the Arabian Peninsula, high levels of cryptic diversity among reptiles, and especially in geckos, have recently been revealed. Mountain ranges within the peninsula were shown to contain the highest richness of reptile endemicity, serving as refugia to species less adapted to the hyper-arid conditions of the lowlands. With...

The European stag beetle Lucanus cervus cervus is the largest European member of the Lucanidae and is characterised by the extremely enlarged mandibles of the male. Some aspects of the morphology of this species have been extremely well studied, while others have been largely neglected. In the present paper we fill some of these gaps by focusing on...

Among geckos, an ideal group to investigate the roles of population dynamics and ecological adaptation in species diversification are semaphore geckos of the genus Pristurus. Numerous ecologically diverse species can be found across Afro-Arabia. On the one hand, strong environmental gradients promoting ecological speciation are present; on the othe...

Mechanical grasping and holding devices depend upon a firm and controlled grip. The possibility to improve this gripping performance is severely limited by the need for miniaturization in many applications, such as robotics, microassembly, or surgery. In this paper, we show how this gripping can be improved in one application (the endoscopic needle...

The stable environment of subterranean realms is characterized by constant darkness, temperature and humidity, and scarcity of resources. This led to similar adaptations in different lineages of animals, such as the reduction of eyes and pigmentation. It is common textbook knowledge that blindness in cave insects is compensated for by transformatio...

Geometric morphometrics (GM) enable the quantification of morphological variation on various scales. Recent technical advances allow analyzing complex three-dimensional shapes also in cases where landmark-based approaches are not appropriate. Pelvic girdle bones (basipterygia) of Sulawesi ricefishes are 3D structures that challenge traditional morp...

Males of the South American spider genus Mecolaesthus Simon, 1893 are characterised by a dorsally ‘inflated’ prosoma or hump. Here we describe the first representatives of the genus from Ecuador. A phylogenetic analysis of molecular sequence data suggests that the new species are part of an Amazonian-West Indian clade. Whether this clade is sister...

Understanding how ecology shapes the evolution of morphological traits is a major goal in organismal biology. By quantifying force of motion, hypotheses on the function of fundamental tasks of animals like feeding can be tested. Ray-finned fishes use various feeding strategies, classified into three main feeding modes: suction, ram and manipulation...

Teleost fishes show an enormous diversity of parental care, ranging from no care to viviparity with maternal provisioning of embryos. External brooders carry their developing eggs attached to their bodies. This requires the formation of novel morphological structures to support attachment. The pelvic brooding ricefish Oryzias eversi evolved such a...

The tracheal system comprises one of the major adaptations of insects towards a terrestrial lifestyle. Many aspects such as the modifications towards wing reduction or a life in an arid climate are still poorly understood. To address these issues, we performed the first three-dimensional morphometric analyses of the tracheal system of a wingless in...

Insect systematics underwent an impressive “evolution” in the last 20 years, especially in terms of methodology, including analyses of very large molecular data sets, but also refined anatomical techniques. Nevertheless, some older phylogenetic concepts were largely confirmed in the “age of phylogenomics”. A remarkable tree of insects was already p...

Evolutionary radiations are one plausible explanation for the rich biodiversity on Earth. Adaptive radiations are the most studied form of evolutionary radiations, and ecological opportunity has been identified as one factor permitting them. Competition among individuals is supposedly highest in populations of conspecifics. Divergent modes of resou...

Respiratory systems are key innovations for the radiation of terrestrial arthropods. It is therefore surprising that there is still a considerable lack of knowledge. In this review of the available information on tracheal systems of hexapods (with a focus on the apterygote lineages Protura, Collembola, Diplura, Archaeognatha and Zygentoma), we summ...

Extant members of the ancient insect order of stoneflies exhibit a disjunct, antitropical distribution, with one major lineage exclusively occurring in the Southern Hemisphere and the other, with few exceptions, on the Northern continents. Here, we address the biogeographic distribution and phylogenetic relationships of stoneflies using a phylogene...

The morphology of the antennal hearts in the head of Phasmatodea and Embioptera was investigated with particular reference to phylogenetically relevant key taxa. The antennal circulatory organs of all examined species have the same basic construction: they consist of antennal vessels that are connected to ampullae located in the head near the anten...

Background Pelvic brooding is a form of uni-parental care, and likely evolved in parallel in two lineages of Sulawesi ricefishes. Contrary to all other ricefishes, females of pelvic brooding species do not deposit eggs at a substrate (transfer brooding), but carry them until the fry hatches. We assume that modifications reducing the costs of egg ca...

Phylogenomics seeks to use next‐generation data to robustly infer an organism's evolutionary history. Yet, the practical caveats of phylogenomics motivate investigation of improved efficiency, particularly when quality of phylogenies are questionable. To achieve improvements, one goal is to maintain or enhance the quality of phylogenetic inference...

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Acoustic communication is enabled by the evolution of specialised hearing and sound producing organs. In this study, we performed a large-scale macroevolutionary study to understand how both hearing and sound production evolved and affected diversification in the insect order Orthoptera, which includes many familiar singing insects, such as cricket...

The genusLasallegriongen. n.is described, and three species,Lasallegrion koebelei(Crawford1912),comb. n.,Lasallegrion virescens(Strand1911),comb. n., andLasallegrion washingtoni(Girault1915),comb. n., are redescribed and transferred to it.Podagrion holbeiniGirault 1923,syn. n., andPodagrion metatarsumGirault 1929,syn. n., are synonymised here withL...

Key changes in ecological niche space are often critical to understanding how lineages diversify during adaptive radiations. However, the converse, or understanding why some lineages are depauperate and relictual, is more challenging, as many factors may constrain niche evolution. In the case of the insect order Grylloblattodea, highly conserved th...

Burmese amber and amber from other periods and regions became a rich source of new extinct insect species and yielded important insights in insect evolution in the dimension of time. Amber fossils have contributed to the understanding of the phylogeny, biology, and biogeography of insects and other groups, and have also gained great importance for...

The systematic position of the Cretaceous genus †Umenocoleus Chen & Tan is re-evaluated. The re-assignment in 2014 back to Coleoptera is rejected based on numerous morphological features incompatible with such a placement. This includes an orthognathous head, multi-segmented antennae, the lack of a cephalic neck region, the exposed cervical membran...

Earwigs are one of the comparatively species‐poor insect orders. Although various aspects of the phylogeny of this lineage are poorly understood, before the present study, there was a general consensus that Dermaptera comprises two major lineages: the paraphyletic Protodermaptera or ‘lower earwigs’ and the monophyletic Epidermaptera or ‘higher earw...

The morphology, musculature, and function of the feeding apparatus of cockroaches is described in detail and compared with other insects with biting and chewing mouthparts. The mouthparts of cockroaches represent, in most cases, the ancestral condition for winged and neopteran insects. Their head capsule is flattened in a posterior-anterior directi...

Structural features and life habits of described species of the extinct †Alienoptera are evaluated based on previously published studies on the group. Head structures and feedings habits are addressed, as are the locomotor organs, especially the wings and adhesive devices. Suggested pollen feeding habits and the possible role as pollinators are dis...

Phasmatodea comprises over 3,000 extant species and stands out as one of the last remaining insect orders for which a robust, higher-level phylogenetic hypothesis is lacking. New research suggests that the extant diversity is the result of a surprisingly recent and rapid radiation that has been difficult to resolve with standard Sanger sequence dat...

Many hemimetabolous insects produce their own cellulase enzymes from the glycoside hydrolase family 9, first observed in termites and cockroaches. Phasmatodea have multiple cellulases, some of which are multifunctional and can degrade xylan or xyloglucan. To discover when these abilities evolved, we identified cellulases from the Polyneoptera sampl...

We describe a new species, Moridilla jobeli sp. nov., belonging to the marine heterobranch group Aeolidioidea. Up to now, it is only recorded from Bunaken National Park, North Sulawesi, Indonesia. A combination of histological, computer tomographic and scanning electron microscopic methods was applied in order to describe and illustrate the anatomy...

Leiodidae are the second largest subterranean radiation of beetles at family rank. To explore morphological trends linked with troglobiontic habits and characters with potential phylogenetic significance, the head of the cave-dwelling species Troglocharinus ferreri (Cholevinae, Leptodirini) was examined in detail. Overall, the general pattern is si...

Phylogenomics seeks to use next-generation data to robustly infer an organism's evolutionary history. Yet, the practical caveats of phylogenomics motivates investigation of improved efficiency, particularly when quality of phylogenies are questionable. To achieve improvements, one goal is to maintain or enhance the quality of phylogenetic inference...

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 ana...

Polyneoptera represents one of the major lineages of winged insects, comprising around 40,000 extant species in 10 traditional orders, including grasshoppers, roaches, and stoneflies. Many important aspects of polyneopteran evolution, such as their phylo-genetic relationships, changes in their external appearance, their habitat preferences, and soc...

Hemipteroid insects (Paraneoptera), with over 10% of all known insect diversity, are a major component of terrestrial and aquatic ecosystems. Previous phylogenetic analyses have not consistently resolved the relationships among major hemipteroid lineages. We provide maximum likelihood-based phylogenomic analyses of a taxonomically comprehensive dat...

While the number of studies on the role of biodiversity on ecosystem functioning is steadily increasing, a key component of biogeochemical cycling in forests, dead wood decay, has been largely neglected. It remains widely unknown whether and how dead wood decay is affected by diversity loss in forests. We studied the hierarchical effects of tree sp...

External and internal thoracic structures of two carabid species (Trechini) were examined and documented with different techniques. The study has a main focus on the eyeless cave-dwelling specialist Sinaphaenops wangorum, but detailed information is also provided for a species occurring in cave entrances. The phylogenetic background of the structur...

Two new species and two new genera ( Kuboesphasma , Minutophasma ) of Mantophasmatodea that occur in the Richtersveld region of South Africa are described. Kuboesphasmacompactumgen. n., sp. n. was found only in a small area near the village of Kuboes, while Minutophasmarichtersveldensegen. n., sp. n. apparently inhabits a larger area in the Richter...

Insects use different parts of their body to hold on to mating partners, catch prey or to defend themselves, in most cases the mouthparts or the legs. However, in 400 million years of evolution1,2, specialized devices were independently acquired in several groups to adopt these tasks, as for instance modified legs in mantids, assassin bugs or stick...

Morphological adaptations of cave-dwelling organisms including different groups of Coleoptera have fascinated researchers since a long time. Nevertheless, very few detailed documentations of the anatomy of cave beetles using modern techniques are available. In this study, we describe and illustrate external and internal cephalic features of free-li...

Orthoptera, or Saltatoria, represents the most diverse insect group among the lower neopterans or Polyneoptera. The present study provides a detailed comparative investigation of the skeletal and muscular thoracic morphology of 23 orthopteran species. For the first time, we investigate unstudied ensiferan key taxa including Gryllacrididae (raspy cr...

Whirligig beetles, which are known for their rapid gliding on the water surface, have evolved a unique locomotor apparatus. External and internal thoracic structures of Orectochilus villosus (Orectochilini) are described in detail and documented with microcomputed tomography, computer-based 3D reconstructions, and scanning electronic microscopy (SE...

Beamtime and resulting SRμCT data are a valuable resource for researchers of a broad scientific community in life sciences. Most research groups, however, are only interested in a specific organ and use only a fraction of their data. The rest of the data usually remains untapped. By using a new collaborative approach, the NOVA project (Network for...

Head structures of Heterogyrus milloti Legros, 1953 are described in detail and documented with different morphological techniques, including μ-computed tomography and computer-based 3D reconstructions. The results are compared with cephalic conditions found in other gyrinid taxa and the observed characters are interpreted and analysed phylogenetic...

The thoraces of males and flightless females of the geometrid winter moth Nyssiodes lefuarius are described, documented in detail, and compared. Morphological, functional, and evolutionary aspects of the female phenotype are discussed. The flightlessness of female N. lefuarius is linked not only with complex modifications of the skeletomuscular str...

The external and internal head morphology including the musculature of the common earwig Forficula auricularia is described in detail. We specified and corrected previous descriptions and provided a detailed documentation. The head of Forficula is characterized by prognathism, generalized mandibles with a mesal cutting edge distally of the mola and...

Schizodactylidae, splay-footed or dune crickets, represents a distinct lineage among the highly diverse orthopteran subgroup Ensifera (crickets, katydids and allies). Only two extant genera belong to the Schizodactylidae: the winged Eurasian genus Schizodactylus, whose ecology and morphology is well documented, and the wingless South African Comicu...

The skeleto-muscular system of the head of the American cockroach Periplaneta americana is described in detail. The results are compared with previous partial descriptions of the cephalic morphology of this species and other dictyopterans. The head of Periplaneta is, as in other cockroaches, mostly characterized by plesiomorphies such as the typica...

Genes acquired by horizontal transfer are increasingly being found in animal genomes. Understanding their origin and evolution requires knowledge about the phylogenetic relationships from both source and recipient organisms. We used RNASeq data and respective assembled transcript libraries to trace the evolutionary history of polygalacturonase (pec...

In a few insect groups, males pierce the female’s integument with their penis during copulation to transfer sperm. This so-called traumatic insemination was previously confirmed for Strepsiptera but only in species with free-living females. The more derived endoparasitic groups (Stylopidia) were suggested to exhibit brood canal mating. Further, it...

A new insect species (†Alienopterus brachyelytrus Bai, Beutel, Klass, Wipfler et Zhang gen. et sp. nov.) of a new order and family is described, based on a single male embedded in Cretaceous Burmese amber (ca. 99 Ma). Unusual characters are shortened forewings combined with fully developed, operational hindwings, similar as in Dermaptera, and speci...

We examine the functional morphology of the mandibular apparatus, including its driving muscles, of the generalist insect Periplaneta americana using a combination of μ-computed tomography and geometrical modelling. Geometrical modelling was used to determine the changes of the mean fibre angle and length in the mandibular adductor muscle over the...

Knowing the functionality and capabilities of masticatory apparatuses is essential for the ecological classification of jawed organisms. Nevertheless insects, especially with their outstanding high species number providing an overwhelming morphological diversity, are notoriously underexplored with respect to maximum bite forces and their dependency...

When disturbed, adults of the Death's-head hawkmoth (Lepidoptera, Sphingidae: Acherontia atropos) produce short squeaks by drawing in and deflating air into and out of the pharynx as a defence mechanism. We took a new look at Prell's hypothesis of a two-phase mechanism by providing new insights into the functional morphology behind the pharyngeal s...

External and internal head structures of Zorotypus weidneri were examined, documented with SEM images and photographs, and reconstructed 3-dimensionally. The results are compared with published results on Z. hubbardi and with conditions found in other hemimetabolan lineages, with a main focus on the polyneopteran orders and on Psocoptera and Thysan...

Insects are the most speciose group of animals, but the phylogenetic relationships of many major lineages remain unresolved. We inferred the phylogeny of insects from 1478 protein-coding genes. Phylogenomic analyses of nucleotide and amino acid sequences, with site-specific nucleotide or domain-specific amino acid substitution models, produced stat...

The present study compares the mitochondrial genomes of five species of the spittlebug tribe Callitettixini (Hemiptera: Cercopoidea: Cercopidae) from eastern Asia. All genomes of the five species sequenced are circular double-stranded DNA molecules and range from 15,222 to 15,637 bp in length. They contain 22 tRNA genes, 13 protein coding genes (PC...

Zoraptera are a cryptic and enigmatic group of insects. The species diversity is lower than in almost all other groups of Hexapoda, but may be distinctly higher than presently known. Several new species were described from different regions recently. The systematic placement was discussed controversially since the group was discovered 100 years ago...

The antennal hearts of the zorapteran Zorotypus hubbardi and of two psocopteran species (Caecilius sp., Embidopsocus sp.) are described in detail and compared to those of other insects. In Zorotypus, the ampullae of this organ are located dorsally of the antennal base. They are attached to the frontal cuticle of the head capsule and laterally suspe...

The relic silverfish Tricholepidion gertschi is the sole extant representative of the family Lepidotrichidae. Its phylogenetic position is of special interest, since it may provide crucial insights into the early phenotypic evolution of the dicondylian insects. However, the phylogenetic position of T. gertschi is unclear. Originally, it was classif...

Mantophasmatodea was described as a new insect order in 2002. Since then, this small group of wingless insects has developed into one of the best investigated insect taxa. Nevertheless, many aspects of mantophasmatodean morphology as well as their evolutionary relationships remain ambiguous. To determine the phylogenetic relationships of Mantophasm...

Grylloblattodea are one of the most unusual groups of insects and the second smallest order. All known extant species are wingless and exhibit a remarkable preference for cold temperatures. Although their morphology was intensively investigated shortly after their discovery, the systematic position has been disputed for a long time. The placement o...

Despite their enormous diversity, the bauplan of adult winged insects (pterygotes) is remarkably conservative since the Lower Devonian: a five-segmented head, a three-segmented thorax with three pairs of walking legs and an eleven-segmented abdomen without any non-sexual appendages [1 • Snodgrass R.E. Principles of Insect Morphology. McGraw-Hill B...

External and internal head structures of last instar (3rd) larvae, 4th day pupae and adults of Chrysomela populi were examined using a combination of traditional and modern morphological techniques, especially mu-computed tomography and computer-based 3D reconstruction. Morphological differences and similarities between the stages were assessed. In...

One of the most ancient ecosystems in the southeastern U.S. is scrub, often associated with ridge systems, mainly in Florida, which are thought to have been used as refugia during the sea level rises of the Pleistocene era. Following recession of the waters these habitats effectively remained islands due partly to unique soil composition as well as...

In Figure 3b, the shadings used to indicate membership in Neoptera and Chiastomyaria were placed incorrectly. The corrected figure is shown here. FIGURE 3. Tree inference from analysis of the morphological and molecular data. a) Consensus tree of the morphological data analyzed with Bayesian inference, ML, maximum parsimony, and parsimony bootstrap...

Extreme elongation of a part of the intromittent organ, the flagellum, has occurred several times in Criocerinae (Chrysomelidae). These leaf beetles have acquired a specialized pocket to store the flagellum in the abdominal cavity, at the same time allowing a quick control of movements of this structure during copulation. We investigated the morpho...

Polyneoptera or “lower Neoptera” are a lineage of moderate diversity, with ca. 36,000 described species and presently ten orders. The monophyly was discussed controversially and is not well supported by morphological data. The reconstruction of interordinal relationships is one of the remaining big challenges in insect systematics. In this contribu...

The first detailed anatomical study of a primary larva of Meloidae is presented. Thereby techniques such as three-dimensional reconstructions, microtome sections, SEM (scanning electronic microscopy) and CLSM (confocal laser scanning microscopy) are applied. The structural features are discussed in the context of phylogeny, but also possible correl...

The earliest branching event in winged insects, one of the core problems regarding early insect evolution, was addressed using characters of the head. The head is arguably one of the most complex body regions in insects and the phylogenetic information content of its features has been demonstrated. In contrast, the wings and other body parts relate...

The relationships of the three major clades of winged insects - Ephemeroptera, Odonata and Neoptera - are still unclear. Many morphologists favor a clade Metapterygota (Odonata+Neoptera), but Chiastomyaria (Ephemeroptera+Neoptera) or Palaeoptera (Ephemeroptera+Odonata) have also been supported in some older and more recent studies.A possible explan...

Published research on the diversity and evolutionary history of Arthropoda sets a high standard for data collection and the integration of novel methods. New phylogenetic estimation algorithms, divergence time approaches, collaborative tools and publishing standards, to name a few, were brought to the broader scientific audience in the context of a...

The larval head of Protanyderus was examined and documented using innovative techniques, with emphasis on internal structures. A chart listing all head muscles of dipteran larvae and other holometabolan groups is presented in the Supporting Information. The results are compared to conditions found in other nematoceran lineages. The larval head of P...

Cover illustration. The cover image shows the head of a larval Protanyderus spec., a representative of the ancestral dipteran Tanyderidae. The image is a sagittally sectioned reconstruction with foregut, muscles and brain. In this study by Wipfler et al. (pp. 968-980) a detailed documentation of the head structures using innovative morphological me...

In this study, we investigated the relationships among insect orders with a main focus on Polyneoptera (lower Neoptera: roaches, mantids, earwigs, grasshoppers, etc.), and Paraneoptera (thrips, lice, bugs in the wide sense). The relationships between and within these groups of insects are difficult to resolve because only few informative molecular...