NEXAM

In an era of rapid biodiversity decline and climate change, understanding the diversity of life has never been more crucial. Arthropods, which include insects, spiders, and crustaceans, make up around 80% of all known animal species. Yet despite their ecological dominance, the internal anatomy and functional morphology of many groups remain poorly understood.

Modern X-ray imaging techniques, particularly at synchrotron facilities, enable non-destructive 3D visualization of millimeter-sized specimens in stunning detail.

NEXAM – Network for X-ray Imaging of Arthropod Morphology is a research network coordinated by the Karlsruhe Institute of Technology (KIT), specifically by researchers at the Institute for Photon Science and Synchrotron Radiation (IPS) and the Laboratory for Applications of Synchrotron Radiation (LAS).

The network brings together scientists from universities, natural history museums, and research institutions who share a common interest in applying advanced synchrotron-based X-ray imaging techniques to study arthropod structure, function, and evolution - from fossil inclusions to living specimens. Methods include high-throughput microtomography, laminography, in vivo imaging, and phase-contrast X-ray microscopy.

NEXAM provides a collaborative platform to employ KIT’s state-of-the-art X-ray imaging infrastructure, including the high-speed UFO-II and multi-scale LAMINO-II stations at the IMAGE beamline at the KIT Light Source, as well as KIT's HIKA station at PETRA III (DESY).

As a global network of biodiversity stakeholders, NEXAM facilitates interdisciplinary exchange, encourages joint research initiatives, and supports access to imaging pipelines tailored to the needs of arthropod research. Future online workshops and collaborative events will further strengthen this growing community and promote knowledge sharing at the interface of imaging, morphology, and biodiversity science.

Interested in becoming a collaborator? Get in touch!

Related projects and initiatives

Antscan (global collaboration)
Virtual Museum (BMFTR ErUM-Pro, 2025 - 2028)
Joint-X (DFG Research Grant, 2023 - 2026)
SMART-Morph (BMFTR ErUM-Pro, 2022 - 2025)
KI-Morph (BMFTR ErUM-Data, 2022 - 2025)

The NEXAM "arthropod team" at KIT: Thomas van de Kamp, Cristina Vasilița, Jenny Hein, Janes Odar & Pauline Pfeiffer (from left to right).

Coordination

Thomas van de Kamp (scientific coordinator)
Tilo Baumbach (head of IPS & director of LAS)

Arthropod team

Beamline team

Computing team

Gallery

NEXAM Collaborators

Jessica Awad, Naturalis Biodiversity Center, Leiden, The Netherlands
Ming Bai, Chinese Academy of Sciences, Beijing, China
Daria Bajerlein, Adam Mickiewicz University, Poznań, Poland
Viktor Baranov, Doñana Biological Station (EBD-CSIC), Seville, Spain
Rolf G. Beutel, Friedrich Schiller University Jena, Germany
Alexander Blanke, University of Bonn, Germany
Brendon E. Boudinot, Senckenberg Research Institute and Natural History Museum, Frankfurt, Germany
Aleš Buček, Biology Centre CAS, České Budějovice, Czechia
Thies H. Büscher, University of Kiel, Germany
Sebastian Büsse, University of Greifswald, Germany
Gabriela P. Camacho, Museum of Zoology of the University of São Paulo, Brazil
Pierfilippo Cerretti, Sapienza University of Rome, Italy
Sándor Csősz, MTA-ELTE-MTM Ecology Research Group, Budapest, Hungary
Evan P. Economo, University of Maryland, College Park, USA & Okinawa Institute of Science and Technology (OIST), Onna-Son, Japan
Erik T. Frank, University of Würzburg, Germany
Heloise Gibb, Deakin University, Melbourne, Australia
Zachary Griebenow, Colorado State University, Fort Collins, USA
Stanislav N. Gorb, Kiel University, Germany
Emily Hartop, NTNU – Norwegian University of Science and Technology, Trondheim, Norway
Francisco Hita Garcia, Museum für Naturkunde, Berlin, Germany
Michael Heethoff, University of Darmstadt, Germany
Annette Herz, Julius Kühn Institute, Dossenheim, Germany
Hendrik Hölscher, Karlsruhe Institute of Technology, Germany
Martin Husemann, State Museum of Natural History Karlsruhe, Germany
Ela Iwaszkiewicz-Eggebrecht, Swedish Museum of Natural History, Stockholm, Sweden
Petr Janšta, Charles University, Prague, Czechia
Niko Kasalo, Ruđer Bošković Institute, Zagreb, Croatia
Julian Katzke, Okinawa Institute of Science and Technology (OIST), Onna-Son, Japan
Hans Klompen, Museum of Biological Diversity, The Ohio State University, Columbus, USA
Ewa Krzemińska, Polish Academy of Sciences, Warsaw, Poland
Wiesław Krzemiński, Polish Academy of Sciences, Warsaw, Poland
Lars Krogmann, State Museum of Natural History Stuttgart, Germany

David Labonte, Imperial College London, UK
Lily Leahy, La Trobe University, Melbourne, Australia
Anderson Lepeco, University of São Paulo, Brazil
Ziv E. Lieberman, UC Davis, USA
Philipp D. Lösel, Australian National University, Canberra, Australia
Yoko Matsumura, University of Vienna, Austria
Arthur Matte, University of Cambridge, UK
Rudolf Meier, Museum für Naturkunde, Berlin, Germany
Nancy F. Mercado Salas, Museum der Natur Hamburg, Germany
István Mikó, University of New Hampshire Collection of Insects and Arthropods, Durham, USA
Christiane Miller, University of Cambridge, UK
Oliver Niehuis, University of Freiburg, Germany
Maëva Orliac, University of Montpellier, France
Michael C. Orr, State Museum of Natural History Stuttgart, Germany, UC Santa Barbara, USA & Chinese Academy of Sciences, Beijing, China
Evgeny Perkovsky, Ukrainian Academy of Sciences, Kyiv, Ukraine
Anika Preuss, Kiel University, Germany
Christian Rabeling, University of Hohenheim, Germany
Adrian Richter, Senckenberg Research Institute and Natural History Museum, Frankfurt, Germany
Alexander Riedel, State Museum of Natural History Karlsruhe, Germany
Fredrik Ronquist, Swedish Museum of Natural History, Stockholm, Sweden
Frederico F. Salles, Entomological Museum, Federal University of Viçosa, Brazil
Achim H. Schwermann, LWL-Museum für Naturkunde, Münster, Germany
Jan Šobotník, Czech University of Life Sciences, Prague, Czechia
Agnieszka Soszyńska, University of Lodz, Poland
Arnold Staniczek, State Museum of Natural History Stuttgart, Germany
Silke Stoll, Museum Natur und Mensch, Freiburg, Germany
Krzysztof Szpila, Nicolaus Copernicus University in Toruń, Poland
Elijah Talamas, Florida Department of Agriculture and Consumer Services, Gainsville, USA
Sergei Tarasov, Finnish Museum of Natural History, Helsinki, Finland
Gabriele Uhl, University of Greifswald, Germany
Noraly M.M.E. van Meer, University of Florida, Gainesville, USA
Peter Vršanský, Slovak Academy of Sciences, Bratislava, Slovakia

Latest Publications

2025

Matte, A., Guénard, B., Azuma, F., Katzke, J., Gautam, S., Hita Garcia, F., van de Kamp, T. & Economo, E.P. (2025). The evolution of cheaper workers facilitated larger societies and accelerated diversification in ants. Science Advances (in press).
van de Kamp, T. & Economo, E.P. (2025): Large-scale library of 3D ant images captured with synchrotron X-ray microtomography. Nature Methods (Research Briefing; in press).
Katzke, J., Hita Garcia, F., Lösel, P.D., Azuma, F., Faragó, T., Aibekova, L., Casadei-Ferreira, A., Gautam, S., Richter, A., Toulkeridou, E., Bremer, S., Hamann, E., Hein, J., Odar, J., Sarkar, C., Zuber, M., Boomsma, J.J., Feitosa, R.M., Schrader, L., Zhang, G., Csősz, S., Dong, M., Evangelista, O., Fischer, G., Fisher, B. L., Florez Fernandez, J., The GAGA Consortium, García, F., Gómez, K., Grasso, D., de Greef, S., Guénard, B., Hawkes, P.G., Johnson, R.A., Keller, R.A., Larsen, R.S., Linksvayer, T.A., Liu, C., Matte, A., Ogasawara, M., Ran, H., Rodriguez, J., Schifani, E., Schultz, T.R., Shik, J.Z., Sosa-Calvo, J., Tong, C., Tozetto, L., Yoon, S., Yoshimura, M., Zhao, J., Baumbach, T., Economo, E.P. & van de Kamp, T. (2025). High-throughput phenomics of global ant biodiversity. Nature Methods (in press).
Boudinot, B.E., Beutel, R.G., Weingardt, M., van de Kamp, T., Hammel, J.U., Li, D., Richter, A. & Wipfler, B. (2025). The insect head rewound: Clarifications to the groundplan of Hexapoda (Pancrustacea). Arthropod Structure and Development 89: 101490. https://doi.org/10.1016/j.asd.2025.101490
Jandausch, K., Straka, J., van de Kamp, T., Stark, H., Beutel, R.G., Niehuis, O. & Pohl, H. (2025). The paragenital organ of Stylopidae (Insecta: Strepsiptera) and the functional incorporation of the secondary larval exuvia. Journal of Morphology 286: e70088. https://doi.org/10.1002/jmor.70088
Oberski, J.T., Griebenow, Z.H., Adams, R.M.M., Andersen, A., Andrade-Silva, J., Barden, P., Borowiec, M., Brady, S., Csősz, S., Dias, A.M., Dias, R.K.S., Feitosa, R.M., Fernandez, F., Casadei-Ferreira, A., Fisher, B.L., General, D.E.M., Gomez, K., Hammel, J.U., Hawkes, P.G., Janda, M., Khalife, A., Ladino, N., Lieberman, Z., Menchetti, M., Prado, L.P., Probst, R.S., Punnath, A., Richter, A., Silva, R.R., Salata, S., Sánchez-Restrepo, A.F., Schifani, E., Schultz, T.R., Sosa-Calvo, J., Tocora, M.C., Ulysséa, M.A., van de Kamp, T., Wang, W., Williams, J., Camacho, G.P. & Boudinot, B.E. (2025). Ant systematics: past, present, and future. Insect Systematics and Diversity 9: ixaf025. https://doi.org/10.1093/isd/ixaf025
Richter, A., Keller, R.A., Hita Garcia, F., van de Kamp, T. & Economo, E.P. (2025). The evolution of novel mouthpart interlocking mechanisms in ants (Hymenoptera: Formicidae). Zoological Journal of the Linnean Society 205: zlaf113. https://doi.org/10.1093/zoolinnean/zlaf113
Chen, Z.-T., Sroka, P., van de Kamp, T. & Staniczek, A. (2025). Revision of the family Peltoperlidae (Insecta: Plecoptera) in mid-Cretaceous Burmese amber, with description of a new genus. Cretaceous Research 177: 106206. https://doi.org/10.1016/j.cretres.2025.106206
Aibekova, L., Richter, A., Beutel, R.G., van de Kamp, T., Economo, E.P., Griebenow, Z. & Boudinot, B.E. (2025). The mesosoma of Protanilla (Leptanillinae) and the groundplan of the Formicidae (Hymenoptera). Journal of Morphology 286: e70064. https://doi.org/10.1002/jmor.70064
Li, D., Richter, A., Olivos-Cisneros, L., van de Kamp, T., Kronauer, D.J.C., Beutel, R.G. & Boudinot, B.E. (2025). The larval morphology of the clonal raider ant, Ooceraea biroi (Forel, 1907) (Dorylinae), with broader implications for the Formicidae and Hymenoptera. Myrmecological News 35: 161–188. https://doi.org/10.25849/myrmecol.news_035:161
Preuss, A., Schwaha, T., Kovalev, A., Ebmer, D., Herzog, I., Lehnert, K., Grass, C., Sandberg, F., Hamann, E., Zuber, M., van de Kamp, T. & Gorb, S.N. (2025). The ectoparasitic seal louse, Echinophthirius horridus, relies on a sealed tracheal system and spiracle-closing apparatus for underwater respiration. Communications Biology 8: 852. https://doi.org/10.1038/s42003-025-08285-4
Bajerlein, D., Kubicka, A.M., Adamski, Z., Hein, J., Zuber, M., Katzke, J. & van de Kamp, T. (2025). Morphology of mandibles of clown beetles (Coleoptera: Histeridae): hidden diversity, sexual dimorphism, and implications for prey range. Zoological Journal of the Linnean Society 204: zlaf038. https://doi.org/10.1093/zoolinnean/zlaf03
Matsumura, Y., Ramezanpour, S., van de Kamp, T., Gorb, S.N., Kovalev, A. & Rajabi, H. (2025). Correlated evolution of elaborate intromission mechanics during copulation between the sexes in leaf beetles. Journal of the Royal Society Interface 22: 20250155. https://doi.org/10.1098/rsif.2025.0155
Hoag, H.A., Raymond, M., Ulmer, J.M., Schwéger, S., van de Kamp, T., Hamann, E., Zuber, M., Werren, J.H., Gaucher, G., Hazen, M. & Mikó, I. (2025). The cranial gland system of Nasonia spp.: a link between chemical ecology, evo-devo, and descriptive taxonomy (Hymenoptera: Chalcidoidea). Journal of Insect Science 25: ieaf034. https://doi.org/10.1093/jisesa/ieaf034
Naredo, J., Brackett, I., Combita-Heredia, J.O., van de Kamp, T., Zuber, M., Hamann, E., Vázquez, M.M. & Klompen, H. (2025). Structure and variability in the female genital atrium of Uropodina (Acari: Parasitiformes). Arthropod Structure & Development 86: 101428. https://doi.org/10.1016/j.asd.2025.101428
Pichler, C.F., Thelen, R., van de Kamp, T. & Hölscher, H. (2025). Friction Coefficient Evolution of Drying Lubricant in the Joints of Beetles by Friction Force Microscopy. Tribology Letters 73: 31. https://doi.org/10.1007/s11249-025-01963-8
Casadei-Ferreira, A., Camacho, G.P., van de Kamp, T., Lattke, J.E., Feitosa, R.M. & Economo, E.P. (2025). Evolution and functional implications of stinger shape in ants. Evolution 79: 80–99. https://doi.org/10.1093/evolut/qpae142

2024

Edel, C., Rühr, P.T., Frenzel, M., van de Kamp, T., Faragó, T., Hammel, J.U., Wilde, F. & Blanke, A. (2024). Bite force transmission and mandible shape in grasshoppers, crickets, and allies is not driven by dietary niches. Evolution 78: 1958–1968. https://doi.org/10.1093/evolut/qpae121
Szpila, K., van de Kamp, T., Sontag, E., Krzemiński, W., Kopeć, K. & Soszyńska, A. (2024). The hidden world of fossil larvae: description and morphological insights of an immature scorpionfly (Mecoptera: Panorpidae) from the Baltic amber. Zoological Journal of the Linnean Society 202(3): 1–11. https://doi.org/10.1093/zoolinnean/zlae009
Krzemińska, E., Soszyńska, A., Kania-Kłosok, I., Skibińska, K., Kopeć, K., van de Kamp, T., Zhang, Q. & Krzemiński, W. (2024). First fossil mountain midges (Diptera: Deuterophlebiidae) and their evolutionary and ecological implications. Scientific Reports 14: 24864. https://doi.org/10.1038/s41598-024-75389-y
Boudinot, B.E., van de Kamp, T., Peters, P. & Knöllinger, K. (2024). Male genitalia reconsidered: a multimodal anatomy of the bullet ant (Paraponera clavata; Hymenoptera: Formicidae). Journal of Morphology 285: e21757. https://doi.org/10.1002/jmor.21757
Meira, O.M., Beutel, R.G., Pohl, H., van de Kamp, T., Almeida, E.A.B. & Boudinot, B.E. (2024). Bee morphology: a skeletomuscular anatomy of Thyreus (Hymenoptera: Apidae). Journal of Morphology 285: e21751.
https://doi.org/10.1002/jmor.21751
Kasalo, N., Husemann, M., van de Kamp, T. & Skejo, J. (2024). Description of Phelene reinschmidti from Ecuador with notes on the subfamily Lophotettiginae (Orthoptera: Tetrigidae). Evolutionary Systematics 8: 119–125. https://doi.org/10.3897/evolsyst.8.124285
Thakur, H., Agarwal, S., Buček, A., Hradecký, J., Sehadová, H., Mathur, V., Togaev, U., van de Kamp, T., Hamann, E., Liu, R.-H., Verma, K. S., Li, H.-F., Sillam-Dussès, D., Engel, M. S. & Šobotník, J. (2024). Defensive glands in Stylotermitidae (Blattodea: Isoptera). Arthropod Structure & Development 79: 101346.
https://doi.org/10.1016/j.asd.2024.101346

2023

Prebus, M., Georgiev, B., van de Kamp, T., Hamann, E., Baker, I. & Rabeling, C. (2023). The rediscovery of the putative ant social parasite Manica parasitica syn. nov. Biology Letters 19: 20230399. https://doi.org/10.1098/rsbl.2023.0399
Griebenow, Z.H., Richter, A., van de Kamp, T., Economo, E.P. & Liebermann, Z.E. (2023). Comparative morphology of male genital skeletomusculature in the Leptanillinae (Hymenoptera: Formicidae), with a standardized muscular terminology. Arthropod Systematics & Phylogeny 81: 945–1018. https://doi.org/10.3897/asp.81.e104810
Spiecker, R., Pfeiffer, P., Biswal, A., Shcherbinin, M., Spieker, M., Zharov, Y., Hessdorfer, H., Hurst, M., Bellucci, V., Faragó, T., Zuber, M., Herz, A., Cecilia, A., Czyzycki, M., Dias, C.S.B., Novikov, D., Krogmann, L., Hamann, E., van de Kamp, T. & Baumbach, T. (2023). Dose-efficient in vivo X-ray phase-contrast imaging at micrometer resolution by Bragg magnifiers. Optica 10: 1633.
https://doi.org/10.1364/OPTICA.500978
Domínguez, E., van de Kamp, T., Mikó, I., Cuezzo, M.G. & Staniczek, A.H. (2023). The function of wing bullae in mayflies (Insecta: Ephemeroptera) reveals new insights into the early evolution of Pterygota. BMC Biology 21: 268. https://doi.org/10.1186/s12915-023-01750-8
Heethoff, M. & van de Kamp, T. (2023). Artificial intelligence in entomology. Entomologie heute 34: 25–38.
Jandausch, K., van de Kamp, T., Beutel, R.G., Niehuis, O. & Pohl, H. (2023). “Stab, chase me, mate with me, seduce me.” How widespread is traumatic insemination in Strepsiptera? Biological Journal of the Linnean Society 140: 206–223. https://doi.org/10.1093/biolinnean/blad046
Klunk, C.L., Argenta, M.A., Rosumek, F.B., Schmelzle, S., van de Kamp, T., Hammel, J.U., Pie, M.R. & Heethoff, M. (2023). Biomechanical performance of ant mandibles under bite-loading demands. Scientific Reports 13: 16833.
https://doi.org/10.1038/s41598-023-43944-8
Moser, M., Ulmer, J.M., van de Kamp, T., Vasilita, C., Renninger, M., Mikó, I. & Krogmann, L. (2023). Surprising morphological diversity in ceraphronid wasps revealed by a distinctive new species of Aphanogmus (Hymenoptera: Ceraphronoidea). European Journal of Taxonomy 864: 146–166. https://doi.org/10.5852/ejt.2023.864.2095
Ulmer, J., Mikó, I, Richter, A., Helms, A., van de Kamp, T. & Krogmann, L. (2023). ‘Social glands’ in parasitoids? – Convergent evolution of metapleural glands in Hymenoptera. Insect Systematics and Diversity 7: 1–10. https://doi.org/10.1093/isd/ixad001

2022

Olmi, M., Capradossi, L., Eggs, B., van de Kamp, T., Perkovsky, E.E., Guglielmino A. & Vasilenko, D.V. (2022). A new species of Bocchus from Upper Eocene Rovno amber (Hymenoptera: Dryinidae). Journal of Hymenoptera Research 92: 257–272. https://doi.org/10.3897/jhr.92.87084
Jandausch, K., Michels, J., Kovalev, A., Gorb, S.N., van de Kamp, T., Beutel, R.G., Niehuis, O. & Pohl, H. (2022). Have female twisted-wing parasites (Strepsiptera) evolved tolerance traits in response to traumatic penetration? PeerJ 10: e13655. https://doi.org/10.7717/peerj.13655
Lieberman, Z., Billen, J., van de Kamp, T. & Boudinot, B. (2022). The ant abdomen: skeletomuscular and soft tissue anatomy of Amblyopone australis (Hymenoptera: Formicidae). Journal of Morphology 283: 693–770. https://doi.org/10.1002/jmor.21471
van de Kamp, T., Mikó, I., Staniczek, A., Eggs, B., Bajerlein, D., Faragó, T., Hagelstein, L., Hamann, E., Spiecker, R., Baumbach, T., Janšta, P. & Krogmann, L. (2022). Evolution of flexible biting in hyperdiverse parasitoid wasps. Proceedings of the Royal Society B 289: 20212086.
https://doi.org/10.1098/rspb.2021.2086