Marie K. Hörnig

Current information on the website of the University of Greifswald

Marie successfully defended her PhD thesis in January 2019.

My main research interest is the evolution of arthropods, with a focus on the evolution of ontogenetic patterns including fossil representatives. During my diploma thesis, I especially investigated the insect group Dictyoptera.


Modern dictyopteran insects, which comprise roaches, termites and mantodeans, show a wide range of social behaviour including different types of brood care: carrying an ootheca ("egg packages"), ovovivipary (retraction of oothecae into brood pouch) right up to vivipary. Different species also feed and protect the young after hatching and exhibit a sub-social and even eu-social behaviour.


Roach-like insects possess a long-ranging fossil record. They already dominated the fauna of Carboniferous forests 300 million years ago.

My research focusses on the evolution of ontogenesis and the deep-time aspects of social and brood care behaviour. Furthermore, I investigated the origin of the different morphotypes within Dictyoptera.


Behaviour cannot directly be observed in extinct species, yet the morphology of fossilised organisms can show some aspects that indicate specific behaviour (e.g. predation) or different grades of intraspecific sociality and brood care. For reconstructing the evolution of these aspects I investigated extinct fossilized representatives of Dictyoptera (especially fossil inclusions in amber) and compared these with observations of extant species.


Different methods are used for the documentation of the fossilised and extant insects, e.g. light microscopy and composite macrophotography under polarised light, stereo photography, micro-computed tomography, and virtual surface reconstructions.


During my PhD studies I broaden this approach and investigate the evolution of reproductive strategies of fossil insects in general.





09/2016: First place for poster presentation. Annual meeting of the German Palaeontological Society, Dresden


09/2015: Third place for poster presentation. Annual meeting of the German Zoological Society, Graz



Fellowships and grants:


05/2015–12/2017: PhD fellowship of the Studienstiftung des Deutschen Volkes


09/2016: travel grant of Glaxo Smith Kline for the Annual meeting of the German Palaeontological Society, Dresden


04/2015: Bogislaw PhD fellowship of the Ernst-Moritz-Arndt-University of Greifswald (declined in favour of Studienstiftung fellowhip)


2014: EU-Synthesys grant for 3 weeks at the Natural History Museum Vienna


2013: EU-Synthesys grant for 2 weeks at the Zoological Museum Copenhagen



Research visits:


07/2015: Natural History Museum Vienna


07/2014: Zoological Museum Copenhagen



Selected publications:

Complete list on the website of the University of Greifswald


2022 / early view


Baranov, V., Pérez-de la Fuente, R., Engel, M.S., Hammel, J.U., Kiesmüller, C., Hörnig, M.K., Pazinato, P.G., Stahlecker, C., Haug, C. & Haug, J.T. 2022. The first adult mantis lacewing from Baltic amber, with an evaluation of the post-Cretaceous loss of morphological diversity of raptorial appendages in Mantispidae. Fossil Record 25, 11–24. Free access PDF Supplement 1 Supplement 2


Haug, J.T., Kiesmüller, C., Haug, G.T., Haug, C. & Hörnig, M.K. early view. A fossil aphidlion preserved together with its prey in 40 million-year-old Baltic amber. Palaeobiodiversity and Palaeoenvironments. Free access PDF


Hörnig, M.K., Haug, C., Müller, P. & Haug, J.T. 2022. Not quite social – possible cases of gregarious behaviour of immatures of various lineages of Insecta preserved in 100-million-year-old amber. Bulletin of Geosciences 97, 69–87. Free access PDF


Kiesmüller, C., Haug, J.T., Müller, P. & Hörnig, M.K. 2022. A case of frozen behaviour: A flat wasp female with a beetle larva in its grasp in 100-million-year-old amber. Fossil Record 25, 287–305. Free access PDF


Kiesmüller, C., Haug, J.T., Müller, P. & Hörnig, M.K. 2022. Debris-carrying behaviour of bark lice immatures preserved in 100 million years old amber. PalZ 96, 231–258. Free access PDF


Zippel, A., Baranov, V.A., Hammel, J.U., Hörnig, M.K., Haug, C. & Haug, J.T. 2022. The first fossil immature of Elmidae: an unusual riffle beetle larva preserved in Baltic amber. PeerJ 10, e13025. Free access PDF


Zippel, A., Haug, C., Gauweiler, J., Hörnig, M.K., Haug, G.T. & Haug, J.T. 2022. A small beetle larva preserved in 23-million-year-old Mexican amber: possible first fossil record of an immature variegated mud-loving beetle. Boletín de la Sociedad Geológica Mexicana 74, A150322. Free access PDF




Baranov, V. A., Engel, M. S., Hammel, J., Hörnig, M. K., van de Kamp, T., Zuber, M. & Haug, J. T. 2021. Synchrotron-radiation computed tomography uncovers ecosystem functions of fly larvae in an Eocene forest. Palaeontologia Electronica 24(1), a07. Free access PDF


Melzer, R. R., Spitzner, F., Šargač, Z., Hörnig, M. K., Krieger, J., Haug, C., Haug, J. T., Kirchhoff, T., Meth, R., Torres, G. & Harzsch, S. 2021. Methods to study organogenesis in decapod crustacean larvae II: analysing cells and tissues. Helgoland Marine Research 75, art. 2. Free access PDF


Schädel, M., Hörnig, M. K., Hyžný, M. & Haug, J. T. 2021. Mass occurrence of small isopodan crustaceans in 100-million-year-old amber: an extraordinary view on behaviour of extinct organisms. PalZ 95, 429–445. Free access PDF


Zippel, A., Kiesmüller, C., Haug, G.T., Müller, P., Weiterschan, T., Haug, C., Hörnig, M.K. & Haug, J.T. 2021. Long-headed predators in Cretaceous amber—fossil findings of an unusual type of lacewing larva. Palaeoentomology 4, 475–498. Free access PDF Suppl Tab 1 Suppl Tab 2

Suppl Tab 3 Suppl Text 1 Suppl Fig 1





Herrera-Flórez, A. F., Braig, F., Haug, C., Neumann, C., Wunderlich, J., Hörnig, M. K. & Haug, J. T. 2020. Identifying the oldest larva of a myrmeleontiformian lacewing – a morphometric approach. Acta Palaeontologica Polonica 65, 235–250. Free access PDF Suppl.


Hörnig, M. K., Kiesmüller, C., Müller, P., Haug, C. & Haug, J. T. 2020. A new glimpse on trophic interactions of 100-million-year old lacewing larvae. Acta Palaeontologica Polonica 65, 777–786. Free access PDF





Hörnig, M. K., Fischer, T. C. & Haug, J. T. 2019. Caught in the act of hatching–a group of heteropteran nymphs escaping from their eggs preserved in Dominican amber. Palaeodiversity 12, 123–134. Free access PDF


Kiesmüller, C., Hörnig, M. K., Leipner, A., Haug, C. & Haug, J. T. 2019. Palaeozoic palaeodictyopteran insect nymphs with prominent ovipositors from a new locality. Bulletin of Geosciences 94, 23–40. Free access PDF





Haug, J. T., Haug, C., Neumann, C., Sombke, A. & Hörnig, M. K. 2018. Early post-embryonic polyxenidan millipedes from Saxonian amber (Eocene). Bulletin of Geosciences 93, 1–11. Free access PDF


Hörnig, M. K., Haug, C., Schneider, J. W. & Haug, J. T. 2018. Evolution of reproductive strategies in dictyopteran insects—clues from ovipositor morphology of extinct roachoids. Acta Palaeontologica Polonica 63, 1–24. Free access PDF





Haug, J. T., Nagler, C., Haug, C. & Hörnig, M. K. 2017. A group of assassin fly pupae preserved in a single piece of Eocene amber. Bulletin of Geosciences 92, 283–295. Free access PDF


Haug, J. T., Poschmann, M., Hörnig, M. K. & Lutz, H. 2017. A crustacean with eumalacostracan affinities from the Early Devonian Hunsrück Slate (SW Germany). Papers in Palaeontology 3, 151–159. DOI 10.1002/spp2.1070


Hörnig, M. K., Haug, J. T. & Haug, C. 2017. An exceptionally preserved 110 million years old praying mantis provides new insights into the predatory behaviour of early mantodeans. PeerJ 5, e3605. Free access PDF


Nagler, C., Hörnig, M. K., Haug, J. T., Noever, C., Høeg, J. T. & Glenner, H. 2017. The bigger, the better? Volume measurements of parasites and hosts: Parasitic barnacles (Cirripedia, Rhizocephala) and their decapod hosts. PLoS ONE 12, e0179958. Free access PDF





Hörnig, M. K., Sombke, A., Haug, C., Harzsch, S. & Haug, J. T. 2016. What nymphal morphology can tell us about parental investment – a group of cockroach hatchlings in Baltic Amber documented by a multi-method approach. Palaeontologia Electronica 19(1), art. 5A, 20 pp. Free access

PDF Suppl.


Liu, Y., Melzer, R. R., Haug, J. T., Haug, C., Briggs, D. E. G., Hörnig, M. K., He, Y.-y. & Hou, X. 2016. Three-dimensionally preserved minute larva of a great-appendage arthropod from the early Cambrian Chengjiang biota. Proceedings of the National Academy of Sciences of the United States of America (PNAS) 113, 5542–5546. DOI 10.1073/pnas.1522899113





Dittmann, I. L., Hörnig, M. K., Haug, J. T. & Haug, C. 2015. Raptoblatta waddingtonae n. gen. et n. sp. – an Early Cretaceous roach-like insect with a mantodean-type raptorial foreleg. Palaeodiversity 8, 103–111. Free access PDF


Haug, J. T., Hädicke, C. W., Haug, C. & Hörnig, M. K. 2015. A possible hatchling of a jumping bristletail in 50 million years old amber. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 278, 191–199. DOI 10.1127/njgpa/2015/0523





Hädicke, C. W., Hörnig, M. K., Haug, C. & Haug, J. T. 2014. New data on fossil Archaeognatha from Baltic amber and the origin of the insect ovipositor. Palaeodiversity 7, 167–183. Free access PDF Suppl.1 Suppl. 2


Hörnig, M. K., Haug, C., Herd, K. J. & Haug, J. T. 2014. New insights into dictyopteran early development: smallest Palaeozoic roachoid nymph found so far. Palaeodiversity 7, 159–165. Free access PDF





Hörnig, M. K., Haug, J. T. & Haug, C. 2013. New details of Santanmantis axelrodi and the evolution of the mantodean morphotype. Palaeodiversity 6, 157–168. Free access PDF