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Research of the Workgroup

Biosystematic Research in the sense of Primary Documentation

Our research is mainly in the field of biosystematics, with emphasis on the documentation of metazoan organisms, especially Arthropoda and Nemathelminthes, furthermore on comparative morphology and ontogeny, and on the development of standards in descriptions and terminology.

Our focus lies on organismic biology (comparative morphology, external morphology, ontogeny, etc.) and documentation. Methods include, as said, the stringent use of the Hennigian and Axian approach of Phylogenetic Systematics, but of course in the sense of advancing and improving this approach rather than being pure followers. This is particularly by accepting fossil evidence.

We study mainly arthropods, particularly crustaceans based partly on extant material and partly on fossils. This is also our specific focus in teaching (generally we considere the entire Metazoa!). For this, we rear some animals or breed larvae, or collaborate with colleagues roviding such material. Special focus is on small crustaceans such as, at present, maxillopods and cephalocarids.

A special topic of our research is the work on the exceptional 'Orsten'-Fossils, mainly arthropods and nemathelminths. Dieter Waloszek started this work 30 years ago, when he received an assistant position in Bonn in the lab of the late Professor Klaus J. Müller. Prof. Müller wasis the discoverer of the Orsten fauna assemblages, which subsequently have become known from several continents. Klaus Müller permitted Dieter Waloszek to continue the reseaerch on this material, which DW transferred to Ulm.

The exceptional material laid the foundation for: a) new standards in the description and illustrative documentation of fossils; b) the interpretation of structures and structural systems, c) the understanding of evolutionary processes and 4) the evolution and phylogeny of particularly the Arthropoda and Crustacea. Lastly all these hardcore data can be used as a testtool against speculative hypotheses about phylogenies or interpretations of features. The material also helped in the improvement of understanding flat-preserved fossil material. Only such well-preserved fossils can yield a reliable data source. When available they are, as once lived animals, teh true source for the reconstruction of evolutionary relationships – and superior to any otehr data, especally the molecular data, which are just claimed to be something valuable but cannot show anything useful – here in the field of phylogenetics, of course. They also cannot help anything in terms of ontogenies anotehr important aspect of evolution. And also here the 'Orsten' fossils provide most beautifully preserved larvae for evolutionary interpretations – being able to serve as the proof of pathways.

Research topics summarized

  • Arthropods and Nemathelminths. Basic problem there: The unclear relationship situation of arthropods with eitehr annelids or nemathelminths, i.e. teh conflicting Ecdysozoa and Articulata Hypotheses. None of them is persuading at present, and Andreas Maas is just about to finish a manuscript on this.

    • Research on arthropods and crustaceans in partiicular: alone or with national and international partnern, e.g. from Copenhagen; this also includes more generalizing papers on structures and evolutionary aspects

    • Reseerach on neozoans: e.g. the common Gammarus roeseli and the invader Dikerogammarus villosus

    • Resaerch on the 'Orsten' of Sweden and similar lagerstätten: 'Orsten' research could be particularly enhanced with the foundation of the international reyearch group C.O.R.E. in 2005, see C.O.R.E. Group website, details, animals, infos.

    • Research on fossils

1. Research on Living Animals

1.1. Arthropod and Particularly Crustacean Limbs

  • The appendages and are the tools of arthropods, the most diversified and species-rich animal taxon on Earth. They and their details are, therefore, of special interest with regard to the biology, fuctional mrophology, evolution and phylogeny of arthropods.

    In our project we focus on crustacean limbs and their evolution, particular from the point of locomotion and feeding. Aims are the understanding of the ontogenetic pathways of particular structures and structural systems around the limbs, the use of limbs and their details for phylogeny and the creation of reliable evolutionary scenarios. Methods apply include SEM work and lightmicroscopy from standard LM to clsm and more techniques, incl. new staining techniques (see also Haug et al. 2009 for a review of new methods applied – click Publications in the row above).

    One paper on the epipodites/gills of crustaceans has been published, aA manuscript reviewing crustacean limbs and their evolution will be published in fall 2010.

Output:

  • Maas, A. Haug, C., Haug, J.T., Olesen, J., Xiguang Zhang & Waloszek, D. 2009. Early Crustacean Evolution and the Appearance of Epipodites/Gills. – Arthropod Systematics & Phylogeny 67(2), 255-273. PDF

1.2. Project on Cephalocarid Crustacea

  • Cephalocarida is a rare taxon of Crusatcea with not more than approx. 10 species currently described. This project is a collaboration with Jørgen Olesen, Copenhagen. It started some years ago and will be finished hopefully soon. It focuses on SEM work on material on a Lightiella species, L. monniotae.

    In our project we consider special aspects of head, trunk and limb morphology, and a few details could be added on immature stages to discuss developmental changes. The new data cast some doubt on the validity of taxonomic discrimination of the Lightiella species.
    A manuscript is currently under construction.

1.3. Project on Mystacocarida and its Relationships to Copepoda and Cambrian 'Orsten' Skaracarida

  • Another collaboration with Jørgen Olesen, Copenhagen. This project will also, hopefully, be finished soon.

1.4. Project on the Dorsal Shield of Crustacea and Arthropoda in General

  • Carolin Haug's PhD work. More soon.

1.5. Project on the Pleon and Uropods of Malacostraca/Eumalacostraca

  • Verena Kutschera's diploma and PhD work. More soon.

Finished: 1996 – 2002: Investigations of cirripede larvae with Dr. J.T. Høeg, Copenhagen and Dr. A. Rybakov, Vladivostok, and on the embryonic development of phyllocarids with Dr. J. Olesen, Copenhagen

Output:

  • Rybakov, A.V., Korn, O.M., Høeg, J.T. & Waloszek, D. 2002: Larval Development in Peltogasterella studied by scanning electron microscopy (Crustacea: Cirripedia: Rhizocephala). Zoologischer Anzeiger 241, 199-221.
  • Olesen, J. & Walossek, D. 2000: Limb ontogeny and trunk segmentation in Nebalia species (Crustacea, Malacostraca, Leptostraca). Zoomorphology 120, 47-64.
  • Walossek, D., Høeg, J.T. & Shirley, T.C. 1996: Larval development of the rhizocephalan cirripede Briarosaccus tenellus (Maxillopoda: Thecostraca) reared in the laboratory: A scanning electron microscopy study. Hydrobiologia 328, 9-47.

1.6. EU-Molmorph Project on the Specific Fate of the 2nd Limb-bearing Head Segment of Arthropoda – From 11/05 – 10/09

  • Within this Eu-funded PhD project, our former doctorate student Yu Liu from Kunming, China – he finished successfully last summer –, investigated the specific fate of the second appendage-bearing body segment of Arthropoda s. str., which became cephalized in Euarthropoda. The appendage of this post-antennular and also first post-oral segment was originally (in Arthropoda s. str.) a simple trunk limb and as serial as all other more posterior appendages; the according segment was likewise "normal-sized". Not only the appendage of the more anterior segment, the antennula (innervated by the deuterocerebrum) suffered from severe modifications later in the evolutionary lineage of Arthropoda s. str. and in the lineages of the different euarthropod taxa, but also the second appendage, originally the first trunk limb and still not cephalized.

    Even more, the whole post-antennular segment, innervated by the tritocerebrum, was apparently affected and, as an extreme, disappeared almost completely together with its limb in adult myriapods and insects. To investigate the detailed changes toward the situation in the different modern taxa, Yu considered both internal (e.g., nervous system, segmental glands, musculature) and external structures (e.g. attachment position of appendages), because they are equally essential for the analysis. Techniques for investigating the neural anatomy have been trained in the University of Copenhagen (Prof. Andreas Wanninger) and RWTH Aachen (Dr. Rudi Loesel).

    Embryonic development of Latrodectus geometricus (Araneae, Arachnida, Chelicerata) and Gryllus assimilis (Orthoptera, Insecta) were in Yu's special focus. L. geometricus possibly reflects a more original situation, the cricket is seen to reflect the highly modified situation in insects. Yet it is assumed that the situation is much more conservative in the early developmental phase hence Yu's attempt to study embryos, which he studied in Japan at the laboratory of Prof. Machida. Flanked by SEM investigations to monitor the topological situation, his study included alpha-tubulin staining techniques marking the nervous system and confocal laserscanning microscopy.

Output:

  • Liu Y., Maas, A. & Waloszek, D. in press. Early embryonic development of the head region of Gryllus assimilis Fabricius, 1775 (Orthoptera, Insecta). Arthropod Structure & Development.
  • Liu, Y., Maas, A. & Waloszek, D. 2009. Embryology of Latrodectus geometricus. – Arthropod Structure & Development 38(5), 401-416. PDF

1.7. Project on the Mouthpart Morphology of Gammarids

  • This project started as a partly EU-funded project in 2005. We investigated invaders of Lake Constance in southwestern Germany, with the specific duty to investigate the gammarid crustacean Dikerogammarus villosus (on right). This invaded is originally from the Ponto-Caspian area, but has spread out all over western and eastern Europe, has reached the river Rhine and further south Lake Constance (German: Bodensee) and recently also lakes in Switzerland and Lake Garda in Italy. Dikerogammarus villosus has apparently extinguished or, at least, threatens established species (e.g. Gammarus roeseli, Gammarus gammarus etc.). This work was mainly done by Gerd Mayer, using SEM. Our collaboration included Gerhard Maier from Ulm, a specialist for limnic animals.

    This work is now continued and expanded by comparative studies on Gammarus roeseli and more species from Germany, Gerd's doctorate work.

Output:

  • Mayer, G., Maier, G., Maas, A. & Waloszek, D. 2009. Mouthpart morphology of Gammarus roeselii compared to a successful invader, Dikerogammarus villosus. Journal of Crustacean Biology 29(2), 161-174. PDF
  • Mayer, G., Maas, A., Waloszek, D. 2008. Mouthparts of the Ponto-Caspian invader Dikerogammarus villosus Sowinsky, 1894 (Gammaridae). Journal Crustacean Biology 28(1), 1-15. PDF

MORE

Projects on the groundpattern of Arthropoda and on the phylogeny of nemathelminth worms are currently under way. They are mainly done by Andreas Maas.

2. Research on Fossil Material

2.1. Project on 3D Preserved Orsten' Crustacea from China, see also here

  • This project is a collaboration with Professor Zhang from Kunming, China, and Prof. Dr. David Siveter, Leicester, UK, on 'Orsten' fossils from the Lower Cambrian of China. It led to the discovery of the earliest eucrustaceans in the fossil record, pushing down all previous thoughts about branchings further down.

    Our first work on the new entomostracan eucrustacean Yicaris dianensis ended in a publication in NATURE recently (click on Publications above, see also the CORE website).

    Our second paper on an early larval stage not belonging to this species with be published June 22 in Current Biology! Its name will be Wujicaris muelleri – after the famous discoverer of the 'Orsten', Klaus Müller, who died this March.

    This so-called metanauplius is approx. 270 µm long. It could be staged by its possession of the two postmandibular limbs, the maxillulae and maxillae, as limb buds. It is as modern as living larvae, having virtually nothing plesiomorphic in its design. Having a caudal shield spine it even resembles extant barnacle larva, but with its forehead spine it looks more like a fish-louse larva.

Output:

  • Zhang Xi-guang, Maas, A., Haug, J.T., Siveter, D.J. & Waloszek, D. 2010. A eucrustacean metanauplius from the Lower Cambrian. Current Biology. PDF click
  • Zhang Xi-guang, Siveter, D., Waloszek, D. & Maas, A. 2007. An epipodite-bearing crown-group crustacean from the Lower Cambrian. Nature 449(7162), 595–598. PDF

2.2. Project on 3D-preserved 'Orsten' Arthropods from Sweden and Nemathelminthes from Australia

  • Joachim Haug, now postdoc assistant (finished February 2009), is working since his dissertation on the ontogeny series of different 'Orsten' taxa, mainly stem-lineage derivatives of the Crustacea. His beautiful 3D models produced with the help of the software BLENDER can be seen on the CORE website. His project on fossil ontogenies is DFG-funded and expires next year. It also includes investigations of material from the Middle Cambrian Burgess Shale.

  • Christopher Castellani has worked in Ulm for one year on a research position funded by EU MolMorph money. His research project on more than 60 new specimens of pentastomids, which we collected in Sweden in 2004, has been finished and a manuscript been submitted to Palaeontographica. See the CORE website.

    Christopher is now a DFG-funded doctorate student in Ulm, focusing now on the "odds and ends" of the 'Orsten' material.

  • Andreas Maas, Joachim Haug and Dieter Waloßek together with different researchers of the CORE Group have just described or are working on various other 'Orsten' forms to complete the knowledge about the fauna, such as a lobopodian, a tardigrade, a nemathelminth larva, two more eucrustaceans, loricate nemathelminth larvae, and more. For details see the according page on the CORE website.

Recent Output:

  • Haug, J.T., Waloszek, D., Haug, C. & Maas, A. 2010. High-level phylogenetic analysis using developmental sequences: The Cambrian Martinssonia elongata, Musacaris gerdgeyeri gen. et sp. nov. and their position in early crustacean evolution. Arthropod Structure & Development 39, 154-173.
  • Maas, A., Waloszek, D., Haug, J.T. & Müller, K.J. 2009. Loricate larvae (Scalidophora) from the Middle Cambrian of Australia. In: Paterson, J.R. & Laurie, J.R. (eds.): Cambro-Ordovician Studies III. Memoirs of the Association of the Australasian Palaeontologists 37, 281-302.
  • Haug, J.T., Maas, A., Waloszek, D., Donoghue, P.C.J. & Bengtson, S. 2009. A new species of Markuelia from the Middle Cambrian of Australia. In: Paterson, J.R. & Laurie, J.R. (eds.): Cambro-Ordovician Studies III. Memoirs of the Association of Australasian Palaeontologists 37, 303-313.
  • Haug, J.T., Maas, A. & Waloszek, D. 2009b. Ontogeny of two Cambrian stem crustaceans, Goticaris longispinosa and Cambropachycope clarksoni. – Palaeontographica A 289(1-3), 1-43.
  • Stein, M., Waloszek, D., Maas, A., Haug. J.T. & Müller, K.J. 2008. Oelandocaris oelandica revisited. Acta Palaeontologica Polonica 53(3), 462-484.
  • Haug, J.T., Maas, A. Waloszek, D., Liu Yu, Haug, C., Castellani, C. & Müller, K.J. 2008. ‘Orsten’ – Cambrian Meiofauna as a modern Ecotype questioning Speculations of a Transition of Pre-Cambrian Ecotypes and Life Strategies far within the Cambrian. Contributions of the Geosciences Union General assembly 2008, Vienna, 13-18. April, Geophysical Research, published abstracts 10.
  • Maas, A., Waloszek, D., Haug, J.T. & Müller, K.J. 2007: A possible larval round worm from the Cambrian 'Orsten' and its bearing on the phylogeny of Cycloneuralia. In: Laurie, J.R. & Paterson, J.R. (eds.): Papers in honour of John Helsby Shergold, 1938-2006. Memoir of the Association of Australasian Palaeontologists 34, 499-519.
  • Maas, A., Waloszek, D., Mayer, G. & Kristensen, R.M. 2007. A Cambrian Micro-lobopodian and the evolution of locomotion and reproduction of Artrhopoda. Chinese Science Bulletin 52(24), 3385-3392.
  • Waloszek, D., Maas, A., Chen Junyuan & Stein, M. 2007. Evolution of cephalic feeding structures and the phylogeny of Arthropoda. Palaeogeography, Palaeoclimatology, Palaeoecology 254, 273–287.

2.3. 3D and 2D Fossils from Devonian Rhynie and Upper Jurassic Solnhofen Fossil Sites

  • Mainly worked up by Carolin and Joachim T. Haug. Collaboration with verious colleagues, such as Stephen Fayers and Hans Kerp as well as colleagues from Stuttgart and amateurs working on Solnhofen Material.
    More soon.

Output:

  • Haug, C., Haug, J.T. & Waloszek, D. 2009. Neues zur Krebsfauna der süddeutschen Plattenkalke. Archaeopteryx 27, 31-37.
  • Haug, J.T., Haug, C., Waloszek, D., Maas, A., Wulf, M. & Schweigert, G. 2009. Development in Mesozoic scyllarids and implications for the evolution of Achelata (Reptantia, Decapoda, Crustacea). Palaeodiversity 2, 97–110.
  • Haug, C., Haug, J.T., Waloszek, D. 2009. Morphology and ontogeny of the Upper Jurassic mantis shrimp Spinosculda ehrlichi n. gen. n. sp. from southern Germany. Palaeodiversity 2, 111–118.
  • Haug, C., Haug, J.T., Waloszek, D., Maas, A., Frattigiani, R. & Liebau, S. 2009. New methods to document fossils from lithographic limestones of southern Germany and Lebanon. Palaeontologia Electronica 12(3); 6T; 12p.http://palaeo-electronica.org/2009_3/193/index.html
  • Haug, J. T., Haug, C., Maas, A. & Waloszek, D. 2009. Mesozoic mantis shrimps. Terra Nostra 2009(3): 46–47. (publ. Abstr. Jahrestag. Pal. Ges. 2009 Bonn)
  • Haug, J.T., Haug, C., Maas, A., Fayers, S.R., Trewin, N.H. & Waloszek, D. 2009a. Simple 3D Images from Fossil and Recent Micromaterial Using Light Microscopy. Journal of Microscopy 233(1), 93-101.

Finished funded projects, see here

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Last revision: June 22, 2010