Subprojects

Basal Hexapods

Basal hexapods are a paraphyletic assemblage of five groups: Protura (coneheads), Collembola (springtails), Diplura (two-pronged bristletails), Archaeognatha (jumping bristletails) and Zygentoma (bristletails, silverfish and firebrats). They are characterized by their primary lack of wings. Many basal hexapods are of great ecological and economic importance. Especially Collembola play a vital role in soil and leaf litter decomposition. Some cosmopolitan species are pests, like Lepisma saccharina (Zygentoma) or the lucerna flea Sminthurus viridis (Collembola). Since representing the earliest splits of hexapod lineages, resolving the relationships among basal hexapods is a major step to understand the evolutionary success of insects. Yet, phylogenetic relationships of Protura, Collembola and Diplura are unclear. Up till now, EST data is sparsely available for these groups. Our aim is to collect specimens of all phylogenetically relevant subgroups to robustly reconstruct the first splitting events within Hexapoda. The subproject "Basal Hexapods" is a crucial milestone on our way to understand insect evolution as a whole.

Coordination:
Alexander Blanke
, Medical and Biological Engineering Research Group, School of Engineering, University of Hull, UK
Karen Meusemann, University of Freiburg, Evolutionary Biology & Ecology, (Biologie I, Zoology)
e-mail: blanke(KITE)uni-bonn.de
e-mail: mail(KITE)karen-meusemann.de

Contributors: Daniela Bartel Oliver Macek Alexander Blanke Alexander Böhm Makiko Fukui Ryuichiro Machida Karen Meusemann Bernhard Misof Yasutaka Nakagaki Günther Pass Malte Petersen Peter Rühr Kaoru Sekiya Nikola Szucsich Gerald Timelthaler Shigekazu Tomizuka Manfred Walzl

Transodonata

Dragonflies and damselflies (Odonata) are a small insect taxon comprising roughly 6,000 species. They are encountered on every continent except Antarctica and are usually the dominant predators at running waters and ponds. Odonates are important indicators of ecosystem quality and play therefore a vital role for urban planning and assessment. The order Odonata is situated at the base of all winged insects (Pterygota). They exhibit a special form of flight (indirect-direct) and a unique mode of indirect sperm transfer. Yet, their phylogenetic relationship to the other two taxa at the pterygote base - mayflies and all other winged insects (Neoptera) - still remains remains unresolved. Additionally, the family relationships inside the order are completely unclear despite sophisticated molecular and morphological analyses of several decades. The suborder Zygoptera (damselflies) splits into 20 putative families whose monophyly is in many cases doubtful at best. Dragonflies (Anisoptera) comprise 11 well supported monophyletic families. However, some interfamiliar relationships are still heavily disputed. Odonates are among the most ancient insects and fossil records of modern families can be traced back into the Mesozoic. A robust and time calibrated phylogenetic reconstruction of the order Odonata will give exceptionally deep insights into the development and timing of evolutionary key features such as insect flight and sperm transfer, which have presumably played a crucial role in the extremely successful radiation of the megadiverse Neoptera.

Coordination:
Alexander Blanke
, Medical and Biological Engineering Research Group, School of Engineering, University of Hull, UK
e-mail: blanke(KITE)uni-bonn.de

Contributors: Alexander Blanke Carola Greve Karl Kjer Ryuichiro Machida Bernhard Misof Oliver Niehuis Jessica Ware

Polyneoptera

Neoptery, the ability to fold back the wings over the abdomen enables pterygote insects to enter substrate or crevices. It is one of the most important apomorphic features that has contributed to the unparalleled success of insects. In the Carboniferous (ca. 300 million years ago) this key innovation has triggered an extremely accelerated radiation of the eleven extant orders of lower neopteran (or polyneopteran) insects. The reconstruction of the relationships is greatly impeded by the very fast succession of branching events. In fact, the phylogenetic affinities among these groups are one of the most obstinate enigmas in insect systematics. So far several molecular and morphological approaches were made and none of them was successful in solving this problem in a satisfying manner. Using a substantial set of EST data we will make a strong contribution to reconstruct the first great radiation in the evolution of pterygote insects.

Coordination:
Benjamin Wipfler
, Institut für Spezielle Zoologie und Evolutionsbiologie mit Phyletischem Museum, FSU Jena, Germany
Sabrina Simon, Wageningen University, Biosystematics, NL Biosystematics Group
e-mail: benjamin.wipfler(KITE)uni-jena.de
e-mail: info(KITE)sabrina-simon.com

Contributors: Thomas Buckley Mari Fujita Karl Kjer Harald Letsch Paschalia Kapli Ryuichiro Machida Yuta Mashimo Ralph Peters Shota Shimizu Kai Schütte Sabrina Simon Toshiki Uchifune Benjamin Wipfler

Dictyoptera

With more than 9,000 species Dictyoptera is one of most species rich groups of lower neopteran insects. It contains heterophagous solitaire roaches, carnivorous mantids and highly organized, multi-caste termites. One of the most important results of systematic entomology within the last years was the recognition of the fact that termites are deeply nested within Blattodea, thus rendering them social roaches. Nevertheless the phylogeny within Dictyoptera is highly unresolved despite the fact that several morphological and molecular studies target this subject. Combined with a substantial morphological data set that will be gathered in the framework of a DFG-project on morphology of lower neopterous insects, the EST data obtained in the 1KITE project will greatly contribute to a deeper understanding of the evolution of this group.

Coordination:
Benjamin Wipfler
, Institut für Spezielle Zoologie und Evolutionsbiologie mit Phyletischem Museum, FSU Jena, Germany
Sabrina Simon, Wageningen University, Biosystematics, NL Biosystematics Group
e-mail: benjamin.wipfler(KITE)uni-jena.de
e-mail: info(KITE)sabrina-simon.com

Contributors: Philipp Erkeling Dominic Evangelista Harald Letsch Mari Fujita Ryuichiro Machida Karen Meusemann Ralph Peters Kai Schütte Sabrina Simon Jessica Ware Benjamin Wipfler

Paraneoptera

Paraneoptera is a megadiverse group of primarily plant-associated hemimetabolous insects and includes many important agricultural pest species (e.g. aphids). The monophyly (inclusion of Zoraptera) is debatable. Likewise the interrelationships of the three major lineages (Thysanoptera, Psocodea and Hemiptera) are not resolved. Paraneoptera have been repeatedly suggested as sistergroup of Holometabola and are therefore crucial for assessing the character polarity within this lineage. The phylogeny of Psocodea, including the traditional orders Psocoptera and Phthiraptera, is one major issue of the subproject. The second key issue is the monophyly of Homoptera and the interrelationships within Hemiptera. These topics will be addressed in our 1KITE subproject using transcriptomes of a representative sample from all major paraneopteran subgroups.

Coordination:
Frank Friedrich
, Biozentrum Grindel and Zoologisches Museum, Universität Hamburg, Germany
e-mail: frank.friedrich(KITE)uni-hamburg.de

Contributors: Rolf Beutel Frank Friedrich Bernhard Misof Ralph Peters Hans Pohl Kazunori Yoshizawa

Hymenoptera

Hymenoptera are a mega-diverse order of endopterygote insects and encompass more than 120,000 described species. Tightly correlated with the huge taxonomical diversity of Hymenoptera is the diversity of life histories of Hymenoptera species. The order contains literally thousands of endo- and ectoparasitoid species that are or can be used for BioControl of insect pests, and the Hymenoptera venoms represent a rich pharmacopeia of potential new drugs. Many phylogenetic relationships within the order Hymenoptera are still not understood (e.g., what is the sister lineage of the famous and medically important Aculeata, i.e., the Hymenoptera, whose ovipositor has been modified to a stinger?). Associated with this lack of phylogenetic resolution is the difficulty to infer, how often and where specifically endoparasitism, phytophagy, and eusociality have evolved in the Hymenoptera.

Coordination:
Oliver Niehuis
Zoologisches Forschungsmuseum A. Koenig, Zentum für Molekulare Biodiversitätsforschung (ZMB)
Ralph Peters, Zoologisches Forschungsmuseum A. Koenig, Section Arthropoda, Bonn, Germany
e-mail: o.niehuis.zfmk(KITE)uni-bonn.de
e-mail: r.peters(KITE)zfmk.de

Contributors: Lars Krogmann Oliver Niehuis Ralph Peters

Neuropterida

Neuropterida comprise the endopterygote insect orders Neuroptera, Megaloptera, and Raphidioptera. Most systematists consider Neuropterida as being a monophyletic group and possible sister of Coleoptera. Neuropterida exhibit numerous plesiomorphic traits and knowlegde of their phylogenetic origin within endopterygote insects will help understanding the evolution of the mega-diverse holometabolous insects. However, molecular analyses have not been able to unequivocally infer the phylogenetic affinities of Neuropterida and to resolve interordinal relationships within Neuropterida. In this project, we aim to solve the above phylogenetic questions by sequencing the transcriptomes of representatives of all extant families of Neuropterida. The obtained data will for the first time allow us to realistically estimate the evolutionary time scale of neuropterid diversification.

Coordination:
Bernhard Misof
, Zoologisches Forschungsmuseum A. Koenig, Zentum für Molekulare Biodiversitätsforschung (ZMB), Bonn, Germany
e-mail: b.misof.zfmk(KITE)uni-bonn.de

Contributors: Ulrike Aspöck Horst Aspöck Rolf Beutel Carola Greve Karl Kjer Xingyue Liu Karen Meusemann Bernhard Misof Oliver Niehuis Ralph Peters Jes Rust Alexandros Vasilikopoulos Torsten Wappler Ding Yang

Coleoptera

With ca. 386,500 described species Coleoptera is by far the largest group of insects. Aside from this unparalleled diversification, beetles are often attractive animals and some groups such as weevils or leaf beetles have a tremendous economic impact. Despite of numerous older and recent phylogenetic studies crucial questions are still open. The interrelationships of the four suborders are discussed very controversially and the relationships of the major lineages of the extremely species rich Polyphaga are still unclear, like the relationships within the very large series Cucujiformia. The early splitting events within Coleoptera, i.e., the relationships of Archostemata (ca. 40 spp.), Myxophaga (ca. 100 spp.), Adephaga (ca. 45,000 spp.) and Polyphaga (ca. 325,000 spp.), and a reconstruction of the basal branching pattern within the megadiverse polyphagan beetles are the primary issues of this subproject. Based on the phylogenetic reconstruction, scenarios will be developed for the evolutionary history of beetles in the Mesozoic.

Coordination:
Adam Slipinski
, CSIRO Ecosystem Sciences, Australian National Insect Collection Acton, Canberra, ACT, Australia
e-mail: adam.slipinski(KITE)csiro.au

Contributors: Rolf Beutel Michael Balke Hermes Escalona Frank Friedrich Harald Letsch Karl Kjer Duanne McKenna José Luis Navarrete-Heredia Ralph Peters Seunggwan Shin Adam Slipinski

Lepidoptera

Butterflies and moths (Lepidoptera), together with Trichoptera, form the endopterygote group Amphiesmenoptera. Lepidoptera have scales on their wings, and this is one of several characters that unites the Order. Larvae are generally terrestrial, and the majority is phytophagous. The pupa is typically created in the ground, within bark, or attached to substrate. Adults of lower Lepidoptera, in the families Micropterigidae, Agathiphagidae, and Heterobathmiidae, are mandibulate. Eriocraniidae have mandibles in the pupal stage, but the adult has a short proboscis. Many of these lower lepidopteran lineages have larvae that are leaf miners or stem borers, internally feeding within plants. More than 98% of all Lepidoptera belong in the large clade Ditrysia. This group has been proposed to have radiated with flowering plants during the Cretaceous. The ditrysian female genital system is very different from lower Lepidoptera, females of Ditrysia have a genital opening for mating, and a second opening for oviposition.

The large monophyletic clade Apoditrysia, characterized by modifications to the second abdominal sternite, consists of most of the larger moths and butterflies, but includes some lineages traditionally placed in the 'Microlepidoptera'. Recent molecular studies have shown that the microlepidoptera are a paraphyletic assemblage. Similarly, the 'Macrolepidoptera', consisting of large moths and butterflies, is also probably paraphyletic. While considerable progress was made in recent years (e.g., Regier et al. 2009; 2013; Mutanen et al. 2010; Cho et al. 2011), the phylogenetic relationships are far from being clear, especially with respect to the backbone of Lepidoptera. A reliable interpretation of character evolution within the group is still needed. This subproject will utilize extensive molecular data to construct a phylogeny of Lepidoptera.

Coordination:
Akito Y. Kawahara
, Florida Museum of Natural History, University of Florida, USA
Bernhard Misof, Zoologisches Forschungsmuseum A. Koenig, Zentum für Molekulare Biodiversitätsforschung (ZMB), Bonn, Germany
Ralph Peters, Zoologisches Forschungsmuseum A. Koenig, Section Arthropoda, Bonn, Germany
e-mail: kawahara(KITE)flmnh.ufl.edu
e-mail: b.misof.zfmk(KITE)uni-bonn.de
e-mail: r.peters(KITE)zfmk.de

Contributors: Akito Y. Kawahara Jesse Breinholt Karl Kjer Karen Meusemann Bernhard Misof Ralph Peters Xin Zhou

Trichoptera

Trichoptera or caddisflies, together with Lepidoptera, form the endopterygote group Amphiesmenoptera. Trichopteran larvae are aquatic, and use silk to construct complex cases, nets and retreats. Two suborders of Trichoptera are widely accepted as monophyletic; Annulipalpia, and Integripalpia. Annulipalpian larvae make a wide variety of retreats that are fixed to the substrate. Integripalpians make portable tube-shaped cases. Between these two monophyletic suborders are five "spicipalpian" families whose larvae exhibit an array of case-making behaviors, from free-living predators to tortoise-like cases, and purse cases. Understanding these behaviors, as well as the ancestral habitats and behaviors of both Amphiesmenoptera and Trichoptera require a well resolved phylogeny of Trichoptera, with particular emphasis on the 5 "spicipalpian" families. Past phylogenetic efforts from multiple genes and morphology have left the position of the spicipalpian families unresolved. We hope to resolve this polytomy with approximately 40 transcriptomes from the 1KITE project. Once the backbone phylogeny has been established, our phylogenetic work can continue, using targeted multiplex sequencing of the more promising genes discovered from the transcriptomes. Morphological data will also be collected for this project. This is important because there has not been an order-wide morphological dataset compiled since the 1996 study of Frania and Wiggins (which included only 70 characters). Thus there does not yet exist a large morphological dataset for Trichoptera that can be compared to the molecular phylogeny, and with which we can postulate morphological and behavioral transitions.

Coordination:
Karl Kjer
, Department of Entomology and Nematology, University of California, Davis, CA, USA
e-mail: kjer(KITE)AESOP.Rutgers.edu

Contributors: Frank Friedrich Paul Frandsen Karl Kjer Martin Kubiak Ralph Peters Xin Zhou

Antliophora

Antliophora comprises the small order Mecoptera, usually characterized by a strongly elongated rostrum, the extremely specialized ectoparasitic Siphonaptera, and the highly derived and mega-diverse Diptera, with many economically and medically important species. The name Antliophora - "pump bearers" - was coined by Willy Hennig. However, the sperm pump was recently discarded as an autapomorphy of the group. Nevertheless, the monophyletic origin of Antliophora, arguably the insect group with the highest degree of accumulated apomorphies, appears well founded.

Even though considerable progress was made in recent years (e.g., Wiegmann et al. 2009; Beutel et al. 2010; Yeates et al. 2011), the phylogenetic relationships are far from being clarified, especially with respect to the position of Boreidae (=Neomecoptera) and Nannochoristidae (=Nannomecoptera), and the basal splitting events within Diptera (e.g., Whiting 2002; Wiegmann et al. 2009; Beutel et al. 2010). A reliable interpretation of the character evolution within the group is still wanting. The reconstruction of a well-founded evolutionary scenario is only possible with a very robust phylogeny on one hand, and detailed morphological and anatomical data on the other. A large morphological data set was already acquired in the first stage of a project on the phylogeny and evolution of Holometabola (see Beutel 2010). More morphological studies will be carried out in this subproject and a current project on Diptera (DFG, BE 1789/6-1), and extensive molecular data will be acquired in the framework of 1KITE. The evaluation will not only lead to a reliable reconstruction of the phylogenetic relationships, but also to a much deeper understanding of evolutionary transformations in different life stages of Antliophora. The diversity of larvae, for instance, ranges from caterpillar-like and ground-dwelling feeding stages with well developed compound eyes to extremely slender aquatic larvae and morphologically extremely simplified maggots. The clarify the evolutionary pathways leading to the great diversity in this and other character systems will be a central aim of this subproject.

Coordination:
Brian Wiegmann
, North Carolina State University, Raleigh, NC, USA
e-mail: bwiegman(KITE)ncsu.edu

Contributors: Keith Bayless Rolf Beutel Frank Friedrich Karen Meusemann Bernhard Misof Ralph Peters Katharina Schneeberg Seunggwan Shin Michelle Trautwein Brian Wiegmann David Yeates

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