Prof. Dr. Steffen Harzsch

Zoological Institute and Museum
Cytology and Evolutionary Biology

Soldmannstr. 23
House 6.1
17489 Greifswald

Phone: +49 (0)3834 420 4124
Fax: +49 (0)3834 420 4067

Research Interests / CV

Curriculum vitae Prof. Dr. Steffen Harzsch

Personal Information:

Date and place of birth: 07.06.1966 in Bielefeld/Westfalen/Germany; Marital status: married, two daughters


Current post

Professor for Cytology and Evolutionary Biology, Zoological Institute and Museum, University of Greifswald (since 1.10.2008)

Address: Soldmannstrasse 23, D - 17487 Greifswald, Germany; Phone: 0049-(0)3834/864124, Fax: (0)3834/864067; email: steffen.harzschuni-greifswaldde


Education and academic degrees                          

  • 2002 - 2006         Heisenberg Fellow (DFG) at the University of Ulm
  • 2001                    Habilitation (venia legendi) and degree of a Private Docent PD in Zoology at the University of Bielefeld
  • 1998-2001           Habilitation Stipend (DFG) Wellesley College, Massachusetts, and University of Ulm
  • 1996-1998           Research Stipend (DFG) Wellesley College, Massachusetts, and University of Ulm
  • 1995                    Dr. rer. nat., Univ. of Bielefeld („summa cum laude“)
  • 1991                    Diploma, Univ. of Bielefeld (profession: Diplombiologe)
  • 1985                    Abitur, Hans Ehrenberg Gymnasium, Bielefeld-Sennestadt


Relevant employments and research stays

  • 10/2008 tenured as W2 position as “Professor für Cytologie und Evolutionsbiologie”, Universität Greifswald
  • 12/2006 -9/2008: employed as group leader for neuroanatomy and morphology at the Department of Evolutionary Neuroethology, Max-Planck-Institute for Chemical Ecology Jena
  • 2/1999 – 12/2006: Guest Scientist with Prof. Dr. H. Wolf, Department of Neurobiology, Univ. of Ulm, and Prof. Dr. D. Waloßek, Section Biosystematic Documentation
  • 5/2005 Guest Scientist with Prof. Dr. B. Battelle, Whitney Laboratory, St. Augustine, Florida
  • 10/1996-12/1996, 4/97-5/97, 10/97-12/97, 11/98-12/98, 11/00, 10/02, 11/03 Guest Scientist with Prof. Dr. B. Beltz, Department of Biological Sciences, Wellesley College, Massachusetts
  • 3/1996-4/1996, 7/96, 3/99-4/99 Guest Scientist at the Biologische Anstalt Helgoland, Dr. K. Anger
  • 7/1995 Guest Scientist at the Institute for Zoophysiology, Rheinische Friedrich-Wilhelms Universität Bonn, Prof. Dr. R. Keller, Prof Dr. H. Dircksen
  • 10/1991-5/1995 Research assistant (Bat IIa/2) at the Department of Neuroanatomy, Faculty of Biology, Univ. of Bielefeld, Prof. Dr. Dr. G. Teuchert-Noodt and Prof. Dr. R. Dawirs
  • 4/1991-5/1991 Job at the Monkswood Experimental Station, Huntingdon, U.K., Freshwater Biological Association, Eastern Rivers Group, Prof. Dr. C. Pinder
  • 5/1988-6/1988, 9/1991 Student Instructor Jobs at the Department of Neuroanatomy, Faculty of Biology, Univ. of Bielefeld, Prof. Dr. Dr. G. Teuchert-Noodt
  • 6/1985-8/1986, 7/88-10/88, 6/89-10/89 Employed at the „Experimentell Ökologisches Laboratorium“ of the Biologische Anstalt Helgoland, Section „Mikrofauna“, Dr. G. Uhlig and „Makrobenthos“, Dr. K. Anger


University Committees, Research Funding Panels, Consulting, Advisory Boards

  • Head of the examination board BSc Biology, University of Greifswald (2020 – present)
  • Head of the examination board MSc Biodiversity and Ecology, University of Greifswald (2017 - 2021)
  • Head of the examination board BSc Human Biology, University of Greifswald (2017 - 2020)
  • Member of the review board (Fachkollegium) 203 Zoology of the Deutsche Forschungsgemeinschaft (2020 – present)
  • Board member of the Wissenschaftliche Kommission Niedersachen for evaluating biological research at the universities of Lower Saxony (2021)
  • Chair, Science Advisory Board “Forschungsstiftung Ostsee” (Stralsund; 2021 – present)


Reviewing for funding agencies

  • Deutsche Forschungsgemeinschaft DFG
  • National Science Foundation USA
  • Portal deutsche Forschungsschiffe
  • The Leverhulme Trust
  • Biotechnology and Biological Sciences Research Council (UK)
  • Fonds zur Förderung der wissenschaftlichen Forschung (FWF) Österreich
  • The Netherlands Organization for Scientific Research
  • The German Israeli Foundation for Scientific Research and Development GIF
  • Deutsche Bundesstiftung Umwelt
  • Evangelisches Studienwerk e.V. Villigst
  • HFSP human frontiers science programme


    Research: emerging research topics

    • Understanding the combined effects of fluctuations of multiple environmental drivers on developmental performance of and organogenesis in marine invertebrate larvae
    • Understanding environmentally induced and stochastic developmental components of phenotypic variation in arthropod nervous systems

    Research: current research topics

    • Functional morphology, development and evolution of crustacean olfactory systems.
    • Transition from sea to land: adaptations of the central olfactory pathway in land hermit crabs
    • “Neurophylogeny”: exploring structure, development and evolution of the central, peripheral and neuromuscular nervous system of Arthropoda for understanding the evolution of nervous systems in general.
    • persistent neurogenesis in the brain of ADULT crustaceans

    Research: past research projects

    • Spatial and temporal patterns of proliferation of neuronal stem cells in the crustacean ventral nerve cord
    • Development of the crustacean visual system and insights into arthropod phylogeny
    • Individually identified neurons and insights into arthropod phylogeny
    • Comparative anatomy of the neuromuscular system of arthropod limbs
    • Structure and development of the nervous system in arrow worms
    • Crustacean metamorphosis: development of the sensory, central nervous, and muscle system in relation to behavioural changes during metamorphic larval development.


    GRANTS (total funding per 2021 ca. 3.343.000 €)

    • 10/2021 Research grant DFG HA 2540/20-1 “Embryonic and larval development of the olfactory system in malacostracan crustaceans: a comparative approach” 202.000 € grant
    • 04/2021 Research grant DFG HA 2540/19-1 “Evolutionary specialization of neuronal core circuits in arthropod olfactory systems: structure and function of convergence/divergence in crustaceans versus insects” 220.000 € grant
    • 04/2021 Research grant in DFG Research Traning Group 2020 RESPONSE, phase 3 “Atlantification of the Arctic: range expansion of a marine invertebrate” 251.000 € grant
    • 04/2018 Research grant in DFG Research Traning Group 2020 RESPONSE, phase 2 “Global ocean change: responses of crab larvae to abiotic stress” 251.000 € grant
    • 04/2015 Research grant in DFG Research Traning Group 2020 RESPONSE, phase 1 “Environmental stress, neurogenesis and colonisation in crustaceans” 251.000 € grant
    • 08/2014 Research grant DFG Ha 2540/15-1 “Lebenslang persistierende Neurogenese im zentralen olfaktorischen Pfad der malacostracen Crustaceen: vergleichende Aspekte” 167.000 € grant
    • 06/2014 Research grant DFG Ha 2540/13-1 “An integrative analysis of olfaction in terrestrial hermit crabs”164.000 € grant
    • 03/2012 Research grant DFG Ha 2540/9-1 “The chemical senses of marine isopod crustaceans”140.000 € grant
    • 03/2012 Research grant DFG Ha 2540/8-2 “Evolution of the olfactory system in Myriapoda”93.000 € grant
    • 03/2011 co-applicant in an „Antrag auf Forschungsgroßgeräte nach Art. 91 b GG (DFG INST 292/103-1 FUGG)“  for a nano computer tomograph (Xradia nano Xct-200); total grant sum 800.000 €;30% user time to my department. 240.000 € grant
    • 03/2010 leading applicant in an „Antrag auf Forschungsgroßgeräte nach Art. 91 b GG (DFG INST 292/103-1 FUGG)“  for a spectral confocal laserscan microscope (Leica SP 5 two photon); total grant sum 750.000 €; 40% user time to my department. 300.000 € grant 
    • 07/2009 Research grant DFG Ha 2540/8-1 “Evolution of the olfactory system in Myriapoda: insights from neuroanatomy and behavior”. 90.000 € grant
    • 08/2009 Research grant in DFG SPP 1147 “Metazoan Deep Phylogeny” (DFG Ha 2540/7-3) “Pinning down the phylogenetic position of Chaetognatha: architecture of their nervous system in comparison to potential outgroups”.82.000 € grant
    • 07/2007 – 06/2009 Research grant in DFG SPP 1147 “Metazoan Deep Phylogeny” (DFG HA 2540/7-2) “Developmental neurobiology of Chaetognatha: impact for their phylogenetic position and the evolution of the bilaterian nervous system.” 86.000 € grant
    • 6/2006 – 10/2009 EU Marie Curie Actions, Human Resources and Mobility Activities “Evolution of animal body plans as inferred by develop-mental biology, morphology, molecular, and palaeontology” (No.: 020542, MOLMORPH); joint proposal led by Prof. A. Wanninger (Copenhagen). Title of my application “Arthropod brain architecture, analysis of proneuronal genes in the developing brain, and arthropod phylogeny”. 182.000 € grant
    • 10/2004 – 9/2006 Research grant in DFG SPP 1147 “Metazoan Deep Phylogeny” (DFG HA 2540/7-1) “Neurophylogeny: architecture and development of the nervous system, individually identifiable neurons and the phylogenetic position of the Chaetognatha.”  110.000 € grant
    • 12/2004 – 12/2006 Research Grant (DFG HA 2540/6-1) „Evolution of neurogenetic patterns in Euarthropoda: do insects and crustaceans share homologous brain neuroblasts?” 21.000 € grant
    • 03/2003 – 12/2006 Research Grant (DFG HA 2540/5-1/3) „Evolution of neurogenetic patterns in Euarthropoda: do crustaceans have a neuronal stem cell similar to Drosophila melanogaster neuroblast 7-3?“ 135.850 € grant
    • 12/2004 -12/2006 Heisenberg Fellowship (DFG HA 2540/3-2) 92.400 € stipend
    • 12/2001-12/2004 Heisenberg Fellowship (DFG HA 2540/3-1) and Research Grant (HA 2549/4-1): „Arthropod evolution: development of the visual system and molecular characterization of neuronal stem cells in representatives of the Crustacea, Hexapoda, „Myriapoda” and Chelicerata.“ 35.700 € grant, 132.600 € stipend
    • 12/1998-11/2001 Habilitation Stipend (DFG HA 2540/2-1/3) and Research Grant (HA 2540/2-2/4): „Ontogeny of crustacean thoracic ganglia and the neuromuscular system in the thoracic limbs: impact on our understanding of arthropod phylogeny.”  25.500 € grant, 48.300 € stipend
    • 8/1996-7/1998 Research Stipend (DFG HA 2540/1): „Mechanisms of embryonic neurogenesis and metamorphic reorganisation of the crustacean central nervous system: model of an alternative developmental strategy to the insects“   23.500 € stipend


    Editorships and membership in editorial boards

    • Member of the Editorial Board of “Arthropod Structure and Development” (Elsevier) since 1/2007
    • Member of the Editorial Board of “Zoology” (Elsevier) since 7/2008
    • Guest Editor “Zoology” Special Issue 2021: “Crustaceans in a Changing World
    • Guest Editor „Arthropod Structure and Development“ Special Issue 2007: „Origin and evolution of arthropod visual systems“ Volume 2
    • Guest Editor „Arthropod Structure and Development“ Special Issue 2006: „Origin and evolution of arthropod visual systems“ Volume 1
    • Guest Editor „Arthropod Structure and Development“ Special Issue 2003: „Structure and development of the arthropod nervous system: an evolutionary perspective“


    Reviewing for international journals

    • Acta Ethologica (Springer)
    • Acta Zoologica (Blackwell)
    • Aquatic Biology (Interscience)
    • Arthropod Systematics and Phylogeny
    • Arthropod Structure and Development (Elsevier)
    • BMC Developmental Biology
    • BMC Evolutionary Biology
    • Biological Bulletin
    • Biological Journal of the LINNEAN SOCIETY
    • Bonner Zoologische Beiträge
    • Cell and Tissue Research (Springer)
    • Crustaceana (Brill)
    • Development, Genes and Evolution (Springer)
    • Developmental Biology (Elsevier)
    • European Journal of Histochemistry
    • Evolution and Development (Blackwell)
    • Frontiers in Zoology (BMC)
    • Frontiers in Physiology
    • General and Comparative Endocrinology (Wiley)
    • Heredity (Nature PG)
    • Hydrobiologia (Blackwell)
    • iLife (Cell press)
    • Invertebrate Development & Reproduction
    • Journal of Cell Biology
    • Journal of Comparative Neurology  (Wiley)
    • Journal of comparative Physiology (Springer)
    • Journal of Crustacean Biology (Allen Press)
    • Journal of Experimental Zoology B: Molecular and Developmental Evolution (Wiley)
    • Journal of Experimental Marine Biology and Ecology (Elsevier)
    • Journal of Microscopy (Elsevier)
    • Journal of Neurobiology (Wiley)
    • Journal of Neuroscience Methods (Wiley)
    • Stem Cells and Development
    • Nature
    • New Zealand Journal of Marine and Freshwater Research (Taylor & Francis)
    • Proceedings of the Royal Society B
    • Zoology (Fischer)
    • Zoomorphology (Springer)


    • Šargač Z, Giménez L, González-Ortegón E, Harzsch S, Tremblay N, Torres G (2022) Quantifying the portfolio of larval responses to salinity and temperature in a coastal-marine invertebrate: a cross population study along the European coast. Marine Biology:
    • Harzsch S, Dircksen H, Hansson BS (2022) Local olfactory interneurons provide the basis for neurochemical regionalization of olfactory glomeruli in crustaceans. Journal of Comparative Neurology 530: 1399-1420.
    • Wacker A, Harzsch S (2021) Crustaceans in a changing world. Zoology:

    • Stein W, Harzsch S (2021) The Neurobiology of Ocean Change - insights from decapod crustaceans. Zoology: 125887.

    • Harzsch S, Krieger J (2021) Genealogical relationships of mushroom bodies, hemiellipsoid bodies, and their afferent pathways in the brains of Pancrustacea: recent progress and open questions. Arthropod Structure & Development 65:101100. Open access.

    • Krieger J, Hörnig MK, Kenning M, Hansson BS, Harzsch S (2021) More than one way to smell ashore - Evolution of the olfactory pathway in terrestrial malacostracan crustaceans. Arthropod Structure & Development 60:101022.

    • Torres G, Melzer R, Spitzner F, Šargač Z, Harzsch S, Gimenez L (2021) Methods to study organogenesis in decapod crustacean larvae I: larval rearing, preparation and fixation. Helgoland Marine Research 75:3

    • Melzer RR, Spitzner F, Šargač Z, Hörnig M, Krieger J, Haug C, Haug JT, 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 75:2

    • Torres G, Charmantier G, Wilcokson D, Harzsch S, Giménez L (2021) Physiological basis of interactive responses to temperature and salinity in coastal-marine invertebrate: implications for responses to warming. Ecology & Evolution 11: 7042-7056
    • Šargač Z, Giménez L, Harzsch S, Krieger J, Fjordside K, Torres G (2021) Contrasting offspring responses to variation in salinity and temperature among coastal populations: a maladaptive ecological surprise? Marine Ecology Progress Series 677:51-65 Open access.
    • Perez Y, Müller CHG, Harzsch S (2021) Chapter 15: Phylum Chaetognatha. In: Invertebrate Zoology: a tree of life approach. B. Schierwater, R. Desalle (eds.), Taylor & Francis; pp. 229 - 250
    • Gimenez L, Exton M, Spitzner F, Meth R, Ecker U, Jungblut S, Harzsch S, Saborowksi R, Torres G (2020) Exploring larval phenology as predictor for range expansion in an invasive species. Ecography 43:1-12. Open Access.

    • Polanska M, Kirchhoff T, Dircksen H, Hansson BS, Harzsch S (2020) Functional morphology of the primary olfactory centres in the brain of the hermit crab Coenobita clypeatus (Anomala, Coenobitidae). Cell and Tissue Research 380:449-467.

    • Krieger J, Hörnig MK, Sandeman RE, Sandeman DC, Harzsch S (2020) Masters of communication: The brain of the banded cleaner shrimp Stenopus hispidus (Olivier, 1811) with an emphasis on sensory processing areas. Journal of Comparative Neurology 528(9): 1561-1587. Open access.

    • Machon J, Krieger J, Zbinden M, Ravaux J, Harzsch S (2020) Exploring brain diversity in crustaceans: sensory systems of deep vent shrimps. Neuroforum  26(2):73-84.

    • Steinhoff P, Uhl G, Harzsch S, Sombke A (2020) Visual pathways in the brain of the jumping spider Marpissa muscosa. Journal of Comparative Neurology. doi: 10.1002/cne.24861.


    Harzsch S & Viertel C (2020). Immunolocalization of neurotransmitters and neuromodulators in the developing crayfish brain. In: Sprecher S. (eds) Brain Development. Methods in Molecular Biology, Vol 2047:271-291. Humana, New York, NY Buy chapter or bookvia

    • Bracke A, Domanska G, Bracke K, Harzsch S, van den Brandt J, Bröker B, et al.
      Obesity Impairs Mobility and Adult Hippocampal Neurogenesis (2019) Journal of Experimental Neuroscience. 13:1-10. Open Access

    • Wittfoth C, Harzsch S, Wolff C, Sombke A (2019) The “amphi”-brains of amphipods: new insights from the neuroanatomy of Parhyale hawaiensis (Dana, 1853). Frontiers in Zoology 16:30. Open Access

    • Machon J, Krieger J, Meth R, Zbinden M, Ravaux J, Montagné N, Chertemps T, Harzsch S (2019) Neuroanatomy of a hydrothermal vent shrimp provides insights into the evolution of crustacean integrative brain centers. eLife 2019;8:e47550. Open Access

    • Spitzner F, Giménez L, Meth R, Harzsch S,Torres G (2019) Unmasking intraspecific variation in offspring responses to multiple environmental drivers. Marine Biology. 166:112

    • Knaden M, Bisch-Knaden S, Linz J, Reinecke A, Krieger J, Erland S, Harzsch S, and Hansson BS (2019) Acetoin, a key odor for resource location in the giant robber crab, Birgus latro. Download pdfvia Journal of Experimental Biology 222(12):jeb202929.

    Müller CHG, Harzsch S & Perez Y (2019) Chaetognatha. In Schmidt-Rhaesa A (ed). Handbook of Zoology. Miscellaneous Invertebrates. Walter De Gruyter, Berlin, Boston. Pp. 163-282

    Jirikowski G, Vogt G, Charmantier-Daures M, Charmantier G, Harzsch S (2019) Chapter 3: Organogenesis. In: K. Anger, S. Harzsch, and M. Thiel, editors. The Natural History of the Crustacea, Volume 7: Developmental Biology and Larval Ecology. Oxford University Press, New York, USA; pp. 79-111

    • Spitzner F, Meth R, Krüger C, Nischik ES, Eiler S, Sombke A, Torres G, Harzsch S. (2018) An atlas of larval organogenesis in the European shore crab Carcinus maenas L. (Decapoda, Brachyura, Portunidae). Frontiers in Zoology 15:27.
    • Wittfoth C, Harzsch S (2018) Adult neurogenesis in the central olfactory pathway of dendrobranchiate and caridean shrimps: new insights into the evolution of the deutocerebral proliferative system in reptant decapods. Developmental Neurobiology 78(8): 757-774.
    • Bracke A, Schäfer S, von Bohlen und Halbach V, Klempin F, Bente K, Bracke K, Staar D, van den Brandt J, Harzsch S, Bader M, Wenzel UO, Peters J, von Bohlen und Halbach, O. (accepted) ATP6AP2 over-expression causes morphological alterations in the hippocampus and in hippocampus-related behaviour. Brain Structure and Function
    • Harzsch S, Krieger J (2018) Crustacean olfactory systems: a comparative review and a crustacean perspective on insect olfactory systems. Progress in Neurobiology 161:23-60. Download via
    • Torres G, Spitzner F, Harzsch S, Gimenez L (2018) Ecological Developmental Biology and global ocean change: brachyuran crustacean larvae as models. In: Perspectives on evolutionary (and) developmental biology. Giuseppe Fusco (ed.); Padova University Press; pp. 283-306.
    • Schmelzle S, Heethoff M, Heuveline V, Lösel P, Becker J, Beckmann F, Schluenzen F, Hammel JU, Kopmann A, Mexner W, Vogelgesang M, Jerome NT, Betz O, Beutel R, Wipfler B, Blanke A, Harzsch S, Hörnig MK, Baumbach T, van de Kamp T (2017). The NOVA project: maximizing beam time efficiency through synergistic analyses of SRμCT data. Proc. SPIE 10391, Developments in X-Ray Tomography XI, 103910P; doi: 10.1117/12.2275959;
    • Meth R, Wittfoth C, Harzsch S (2017)Brain architecture of the Pacific White Shrimp Penaeus vannamei  Boone, 1931 (Malacostraca, Dendrobranchiata): correspondence of brain structure and sensory input? Cell and Tissue Research 369(2): 255–271.
    • Steinhoff POM*, Sombke A*, Liedtke J, Schneider JM, Harzsch S, Uhl G. (2017) The synganglion of the jumping spider Marpissa muscosa (Arachnida: Salticidae): insights from histology, immunohistochemistry and microCT analysis. Arthropod Structure & Development 46(2): 156-170. *shared first authorship
    • Audo D, Haug JT, Haug C, Charbonnier S, Schweigert G, Müller CHG, Harzsch S (2016) On the sighted ancestry of blindness - exceptionally preserved eyes of Mesozoic polychelidan lobsters. Zoological Letters 2:13.
    • Harzsch S, von Bohlen und Hallbach O. (2016) A possible role for the immune system in adult neurogenesis: new insights from an invertebrate model. Zoology 119(2):153-157.
    • Hörnig MK, Sombke A, Haug C, Harzsch S, Haug JT (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 19.1.6A.
    • Krieger J, Drew MM, Hansson BS, Harzsch S (2016) Notes on the foraging strategies of the Giant Robber Crab Birgus latro (Anomala) on Christmas Island: Evidence for active predation on Red Crabs Gecarcoidea natalis (Brachyura). Zoological Studies 55:6.
    • Harzsch S, Perez Y, Müller CHG (2016) Chaetognatha. In: Schmidt-Rhaesa A, Harzsch S, Purschke G. (Eds.). Structure and Evolution of Invertebrate Nervous Systems. Oxford University Press, pp. 652-664.
    • Batelle BA, Sombke A, Harzsch S (2016) Xiphosura. In. Schmidt-Rhaesa A, Harzsch S, Purschke G. (Eds.) Structure and Evolution of Invertebrate Nervous Systems. Oxford University Press, pp. 428-442.
    • Stemme T, Harzsch S (2016) Remipedia. In: A. Schmidt-Rhaesa, S. Harzsch, G. Purschke (Eds.). Structure and Evolution of Invertebrate nervous Systems. , Oxford University Press, pp. 522-528.
    • Lehmann D, Melzer RR, Hörnig MK, Michalik P, Sombke A, Harzsch S (2016) Arachnida - excl. Scorpiones. In: A. Schmidt-Rhaesa, S. Harzsch, G. Purschke (Eds.). Structure and Evolution of Invertebrate nervous Systems. , Oxford University Press, pp. 453-477.
    • Krieger J*, Braun P*, Rivera NT, Schubart CD, Müller CHG, Harzsch S (2015) Comparative analyses of olfactory systems in terrestrial crabs (Brachyura): Evidence for aerial olfaction?. PeerJ 3: e1433 *shared first authorship
    • Tuchina O, Koczan S, Harzsch S, Rybak J, Wolff G, Strausfeld NJ, Hansson BS (2015) Central projections of antennular chemosensory and mechanosensory afferents in the brain of the terrestrial hermit crab (Coenobita clypeatus; Coenobitidae, Anomura). Frontiers in Neuroanatomy 9:94.
    • Kenning M, Lehmann P, Lindström M, Harzsch S (2015) Heading which way? Y-maze chemical assays – not all crustaceans are alike. Helogoland Marine Research. Download via
    • Grunow B, Kirchhoff T, Lange T, Moritz T, Harzsch S (in press) Histochemistry on vibratome sections of fish tissue: a comparison of fixation and embedding methods. Aquatic Biology. doi 10.3354/ab00624.
    • Sombke A, Harzsch S (2015) Immunolocalization of histamine in the optic neuropils of Scutigera coleoptrata (Myriapoda: Chilopoda) reveals the basal organization of visual systems in Mandibulata. Neuroscience Letters 594: 111-116.
    • Sombke A, Lipke E, Michalik P, Uhl G, Harzsch S (2015) Potential and limitations of X-ray micro-computed tomography in arthropod neuroanatomy – a methodological and comparative survey. The Journal of Comparative Neurology 523(8): 1281-1295. Open Access
    • Kress T, Harzsch S, Dircksen H (2015) Neuroanatomy of the optic ganglia and central brain of the water flea Daphnia magna (Crustacea, Cladocera). Cell and Tissue Research: DOI 10.1007/s00441-015-2279-4.
    • Harzsch S , Krieger J, Faulkes Z (2015) Astacida. In: A Wanninger (ed.). Evolutionary Developmental Biology of Invertebrates, Springer, pp: 101-152.
    • Harzsch S, Müller CHG, Perez Y (2015) Chaetognatha. In: A Wanninger (ed.). Evolutionary Developmental Biology of Invertebrates, Springer, pp: 215-240.
    • Ito K, Shinomiya K, Ito M, Armstrong JD, Boyan G, Hartenstein V, Harzsch S, Heisenberg M, Homberg U, Jenett A, Keshishian H, Restifo LL, Rössler W, Simpson JH, Strausfeld NJ, Strauss R, Vosshall LB, Insect Brain Name Working Group (2014) A Systematic Nomenclature for the Insect Brain. Neuron 81(4): 755-765.
    • Müller CHG; Rieger V, Perez Y & Harzsch S (2014). Immunohistochemical and ultrastructural studies on ciliary sense organs of arrow worms (Chaetognatha). Zoomorphology 133:167-189.
    • Perez Y, Müller C, Harzsch S (2014) The Chaetognatha: an anarchistic taxon between Protostomia and Deuterostomiea. In: Wägele JW, Bartholomaeus TW, Misof B (Eds.). Deep metazoan phylogeny: The backbone of the tree of life: New insights from analyses of molecules, morphology, and theory of data analysis. pp: 49-78, Walter de Gruyter, Berlin.
    • Sandeman D, Kenning M, Harzsch S (2014) Adaptive trends in malacostracan brain form and function related to behaviour. In: Derby C and Thiel M (Eds.) Crustacean Nervous System and their Control of Behaviour. The Natural History of the Crustacea. Vol. 3. pp: 11-48, Oxford University Press.
    • Kenning M, Harzsch S (2013) Brain anatomy of the marine isopod Saduria entomon Linnaeus, 1758 (Valvifera, Isopoda) with special emphasis on the olfactory pathway. Frontiers in Neuroanatomy. Open Access
    • Zieger E, Bräunig P, Harzsch S (2013): A developmental study of serotonin-immunoreactive neurons in the embryonic brain of the Marbled Crayfish and the Migratory Locust: evidence for a homologous protocerebral group of neurons. Arthropod Structure & Development. 42(6): 507-520.
    • Stemme T, Iliffe T, von Reumont BM, Koenemann S, Harzsch S, Bicker G (in press) Serotonin-immunoreactive neurons in the ventral nerve cord of Remipedia (Crustacea): Support for a sister group relationship of Remipedia and Hexapoda? BMC Evolutionary Biology 13:119
    • Perez Y, Rieger V, Martin E, Müller C, Harzsch S (2013) Neurogenesis in an early protostome relative: progenitor cells in the ventral nerve centre of chaetognath hatchlings are arranged in a highly organized geometrical pattern. Journal of Experimental Zoology Part B: Molecular and Developmental Evolution. 320(3):179-193.
    • Kenning M, Müller C, Wirkner CS, Harzsch S (2013) The Malacostraca (Crustacea) from a neurophylogenetic perspective: new insights from brain architecture in Nebalia herbstii Leach, 1814 (Leptostraca, Phyllocarida). Zoologischer Anzeiger 252: 319-336.
    • Loesel R, Wolf H, Kenning M, Harzsch S, Sombke A (2013) Architectural principles and evolution of the arthropod central nervous system. In: Minelli A, Boxshall G, Fusco G (Eds.) Arthropod Biology and Evolution: Molecules, Development, Morphology. Springer, Berlin
    • Kapp H, Müller C, Harzsch S (2013). Chaetognatha. In: "Spezielle Zoologie. Teil 1: Einzeller und Wirbellose Tiere". Westheide W, Rieger G (eds.). Springer Spektrum Verlag, pp. 812-817.
    • Krieger J, Grandy R, Drew MM, Erland S, Stensmyr MC, Harzsch S, Hansson BS (2012) Giant robber crabs monitored from space: GPS-based telemetric studies on Christmas Island (Indian Ocean). PLoS ONE 7(11): e49809. Open Access
    • Polanska M, Tuchina O, Agricola H, Hansson BS, Harzsch S (2012) Neuropeptide complexity in the crustacean central olfactory pathway: immunolocalization of A-type allatostatins and RFamide-like peptides in the brain of a terrestrial hermit crab. Molecular Brain. 5:29.
    • Stemme T, Iliffe TM, Bicker G, Harzsch S, Koenemann S (2012) Serotonin immunoreactive interneurons in the brain of the Remipedia: new insights into the phylogenetic affinities of an enigmatic crustacean taxon. BMC Evolutionary Biology. 12:168. doi:10.1186/1471-2148-12-168.
    • Krieger J, Sombke A, Seefluth F, Kenning M, Hansson BS, Harzsch S (2012) Comparative brain architecture of the European shore Crab Carcinus maenas (Brachyura) and the Common hermit Crab Pagurus bernhardus (Anomura). Cell and Tissue Research. 348(1):47-69. Download via
    • Wolff G, Harzsch S, Hansson BS, Brown S, Strausfeld NJ (2012) Neuronal organization of the hemiellipsoid body of the land hermit crab Coenobita clypeatus: correspondence with the mushroom body ground pattern. Journal of Comparative Neurology 520: 2824-2846.
    • Wolf H, Harzsch S (2012) Serotonin-immunoreactive neurons in scorpion pectine neuropils: similarities to insect and crustacean primary olfactory centres? Zoology 115: 151-159.
    • Sombke A, Lipke E, Kenning M, Müller CHG, Hansson BS, Harzsch S. (2012) Comparative analysis of deutocerebral neuropils in Chilopoda (Myriapoda): implications for the evolution of the arthropod olfactory system and support for the Mandibulata concept. BMC Neuroscience 13:1.
    • Sintoni S, Benton JL, Beltz BS, Hansson BS, Harzsch S (2012) Neurogenesis in the central olfactory pathway of adult decapod crustaceans: development of the neurogenic niche in the brains of Procambarid Crayfish. Neural Development 7: 1.
    • Sandeman D, Kenning M, Harzsch S. (accepted pending revisions). Adaptive trends in malacostracan brain form and function related to behaviour. In: Derby C and Thiel M (Eds.) Crustacean Nervous System and their Control of Behaviour. The Natural History of the Crustacea. Vol. 3.
    • Harzsch S, Sandeman D, Chaigneau J (2012) Morphology and development of the central nervous system. In: Forest J and von Vaupel Klein JC (Eds.). Treatise on Zoology – Anatomy, Taxonomy, Biology. The Crustacea Vol. 3. Brill, Leiden pp. 9-236.
    • Rieger V, Perez Y, Müller CHG, Lacalli T, Hansson BS, Harzsch, S (2011) Development of the nervous system in hatchlings of Spadella cephaloptera (Chaetognatha), and implications for nervous system evolution in Bilateria. Development, Growth and Differentiation 53:740-759.
    • Missbach C, Harzsch S, Hansson B (2011) New insights into an ancient insect nose: the olfactory pathway of Lepismachilis y-signata (Archaeognatha: Machilidae). Arthropod Structure & Development 40:317-333.
    • Harzsch S, Rieger V, Krieger J, Seefluth F, Strausfeld NJ, Hansson BS (2011) Transition from marine to terrestrial ecologies: Changes in olfactory and tritocerebral neuropils in land-living isopods. Arthropod Strcuture and Development 40: 244-257.
    • Sombke A, Harzsch S, Hansson BS (2011): Organization of deutocerebral neuropils and olfactory behavior in the centipede Scutigera coleoptrata (Linnaeus, 1758) (Myriapoda: Chilopoda). Chemical Senses 36: 43-61.
    • Hansson BS, Harzsch S, Knaden M & Stensmyr M (2011) The neural and behavioral basis of chemical communication in terrestrial crustaceans. In: Chemical communication in Crustaceans., Breithaupt T., Thiel M. (eds.), Springer, 201: 149-174.
    • Stemme T, Bicker G, Harzsch S, Koenemann S (2010) Die rätselhaften Grottenkrebse der Blue Holes. Sind Remipedia primitive Crustaceen oder schwimmende Insekten? Naturhistorica 152: 7-28.
    • Krieger J, Sandeman RE, Sandeman DC, Hansson BS, Harzsch S (2010) Brain architecture of the largest living land arthropod, the Giant Robber Crab Birgus latro (Crustacea, Anomura, Coenobitidae): evidence for a prominent central olfactory pathway? Frontiers in Zoology. 7:25
    • Richter S, Loesel R, Purschke G, Schmidt-Rhaesa A, Scholtz G, Stach T, Vogt L, Wanninger A, Brenneis G, Döring C, Faller S, Fritsch M, Grobe P, Heuer CM, Kaul S, Møller OS, Müller CHG, Rieger V, Rothe BH, Stegner MEJ, Harzsch S (2010). Invertebrate neurophylogeny - suggested terms and definitions for a neuroanatomical glossary. Frontiers in Zoology 7:29.
    • Harzsch S, Kreisl S (2010) Myogenesis in the thoracic limbs of the American lobster. Arthropod Structure & Development 39: 423-435.
    • Drew MM, Harzsch S, Stensmyr M, Erland S, Hansson BS (2010) A review of the biology and ecology of the Robber Crab, Birgus latro (Linnaeus, 1767) (Anomura: Coenobitidae). Zoologischer Anzeiger 249: 45-67.
    • Rieger V, Perez Y, Müller CHG, Lipke E, Sombke A, Hansson BS, Harzsch S (2010) Immunohistochemical analysis and 3D reconstruction of the cephalic nervous system in Chaetognatha: insights into an early bilaterian brain? Invertebrate Biology 129(1): 77-104.
    2005 - 2009
    • Sombke A, Harzsch S, Hansson BS (2009): Brain structure of Scutigera coleoptrata: New insights into the evolution of mandibulate olfactory centers. - Soil Organisms 81 (3): 319-325.
    • Harzsch S, Beltz B, Dircksen H (2009) Development of pigment-dispersing hormone-immunoreactive neurons in the American lobster: possible homologies with the insect circadian pacemaker system. Cell & Tissue Research 335: 417-429
    • Harzsch S, Rieger V, Müller CHG, Perez Y, Sintoni S, Sardet C, Hansson B (2009) Fine structure of the ventral nerve centre and interspecific identification of individual neurons in the enigmatic Chaetognatha. Zoomorphology 128: 53-73
    • Vilpoux K, Bisch-Knaden S, Harzsch S (2008) Mapping of Engrailed-expressing cells in the embryonic central nervous system of the marbled crayfish (Marmorkrebs). Invertebrate Neuroscience: DOI 10.1007/s10158-008-0081-7
    • Harzsch S, Hansson B (2008) Brain architecture in the terrestrial hermit crab Coenobita clypeatus (Anomura, Coenobitidae): neuroanatomical evidence for a superb aerial sense of smell. BMC Neuroscience 9: 1-35
    • Kreissl S, Uber A, Harzsch S (2008) Muscle pioneer cells in the developing limbs of two isopods (Crustacea, Peracarida): an immunohistochemical study using a novel monoclonal antibody against myosin heavy chain. Dev. Genes Evol. 218: 253-265
    • Mayer G, Harzsch S (2008) Distribution of 5-HT-like immunoreactivity in the trunk of Metaperipatus blainvillei (Onychophora, Peripatopsidae): implications for nervous system evolution in Arthropoda. J. Comp. Neurol. 507: 1196-1208
    • Rieger V, Harzsch S (2008) Embryonic development of the histaminergic system in the ventral nerve cord of the Marbled Crayfish (Marmorkrebs). Tissue and Cell 39: 113-126
    • Polanska M, Yasuda A, Harzsch S (2007) Immunolocalisation of crustacean-SIFamide in the median brain and eyestalk neuropils of the marbled crayfish. Cell & Tissue Res. 330: 331-344
    • Sintoni S, Fabritius-Vilpoux K, Harzsch S (2007) The Engrailed-expressing secondary head spots in the embryonic crayfish brain: examples for a group of homologous neurons in Crustacea and Hexapoda? Dev Genes Evol 217: 791-799
    • Mayer G, Harzsch S (2007) Immunolocalization of serotonin in Onychophora argues against segmental ganglia being an ancestral feature of arthropods. BMC Evolutionary Biology 7: 118
    • Harzsch S, Müller CHG (2007) A new look at the ventral nerve centre of Sagitta: implications for the phylogenetic position of Chaetognatha (arrow worms) and the evolution of the bilaterian nervous system. Frontiers in Zoology 4: 14
    • Harzsch S, Melzer RR, Müller CHG (2007) Mechanisms of eye development and evolution of the arthropod visual system: the lateral eyes of Myriapoda are not modified insect ommatidia. Org. Divers. Evol. 7: 20-32
    • Harzsch S, Melzer RR, Müller CHG (2006) Eye development in Myriapoda: implications for arthropod phylogeny. Norw. J. Entomol. 53: 187-190
    • Harzsch S, Vilpoux K, Blackburn DC, Platchetzki D, Brown NL, Melzer R, Battelle BA (2006) Development of the eyes and central visual pathway of the horseshoe crab Limulus polyphemus Linnaeus, 1758 (Chelicerata, Xiphosura). Developmental Dynamics 235: 2641-2655
    • Vilpoux K, Sandeman R, Harzsch S (2006) Early embryonic development of the central nervous system in the Australian crayfish and the Marbled crayfish (Marmorkrebs). Dev. Genes Evol. 216: 209-223
    • Fanenbruck M, Harzsch S (2005) A brain atlas of Godzilliognomus frondosus Yager, 1989 (Remipedia, Godzilliidae) and comparison with the brain of Speleonectes tulumensis Yager, 1987 (Remipedia, Speleonectidae): implications for arthropod relationships. Arthropod Struct. Dev.: 343-378
    • Harzsch S, Wildt M, Battelle B, Waloszek D (2005) Immunohistochemical localization of neurotransmitters in the nervous system of larval Limulus polyphemus Linnaeus, 1758 (Chelicerata, Xiphosura): evidence for a conserved architecture of the protocerebrum in Euarthropoda. Arthropod. Struct. Dev.: 327-342
    • Seitz R, Vilpoux K, Hopp U, Harzsch S, Maier G (2005) Ontogeny of the Marmorkrebs (marbled crayfish): a parthenogenetic crayfish with unknown origin and phylogenetic position. J. Exp. Zool. 303A: 393-405
    • Harzsch S, Wanninger A (2009) Evolution of invertebrate nervous systems: the Chaetognatha as a case study. Acta Zoologica 91: 35-41.
    • Harzsch S (2007) Architecture of the nervous system as a character for phylogenetic reconstructions: examples from the Arthropoda. Species, Phylogeny & Evolution 1: 33-57.
    • Stavenga DG, Harzsch S (2007) Origin and evolution of arthropod visual systems: introduction to Part II. Arthropod Struct. Dev.36: 371-372.
    • Harzsch S, Melzer RR (2006) Editorial: origin and evolution of arthropod visual systems. Arthropod Struct. Dev. 35: 209-210.
    • Harzsch S, Hafner G (2006) Evolution of eye development in arthropods: phylogenetic implications. Arthropod Struct. Dev. 35: 319-340.
    • Harzsch S (2006) Neurophylogeny: architecture of the nervous system and a fresh view on arthropod phyologeny. Comp. Integr. Biol. 46: 162-194.
    • Harzsch S (2005) Bookreview of “Evolutionary Developmental Biology of Crustacea”, Crustacean Issue, edited by G. Scholtz. J. Zool. Systemat. Evol. Res. 43: 344-345.
    • Harzsch S, Müller CHG, Wolf H (2005) From variable to constant cell numbers: cellular characteristics of the arthropod nervous system argue against a sister-group relationship of Chelicerata and “Myriapoda” bur favour the Mandibulata concept. Dev. Genes. Evol. 215: 53-68.
    2000 - 2004
    • Harzsch S (2004) Phylogenetic comparison of serotonin-immunoreactive neurons in representatives of the Chilopoda, Diplopoda and Chelicerata: implications for arthropod relationships. J. Morphol. 259: 198-213
    • Fanenbruck M, Harzsch S, Wägele W (2004) The brain of the Remipedia (Crustacea) and an alternative hypothesis on their phylogenetic relationships. Proceedings of the National Academy of Sciences USA 101: 3868-3873
    • Harzsch S (2003) Evolution of identified arthropod neurons: the serotonergic system in relation to engrailed-expressing cells in the embryonic ventral nerve cord of the American lobster Homarus americanus Milne Edwards, 1873 (Malacostraca, Pleocyemata, Homarida). Dev. Biol. 258:44-56
    • Harzsch S (2002) Phylogenetic significance of the crustacean optic neuropils and chiasmata: a re-examination. J. Comp. Neurol. 453: 10-21
    • Harzsch S, Glötzner J (2002) An immunohistochemical study on structure and development of the nervous system in the brine shrimp Artemia salina Linnaeus, 1758 (Branchiopoda, Anostraca) with remarks on the evolution of the arthropod brain. Arthropod. Struct. Dev. 30: 251-270.
    • Wildt M, Harzsch S (2002) A new look at an old visual system: structure and development of the compound eyes and optic ganglia of the brine shrimp Artemia salina Linnaeus, 1758 (Branchiopoda, Anostraca). J. Neurobiol. 52: 117-132.
    • Wolf H, Harzsch S (2002) Evolution of the arthropod neuromuscular system: 1. Arrangement of muscles and excitatory innervation in the walking legs of the scorpion Vaejovis spinigerus (Wood, 1863) (Vaejovidae, Scorpiones, Arachnida). Arthropod Struct. Dev. 31: 185-202.
    • Wolf H, Harzsch S (2002) Evolution of the arthropod neuromuscular system. 2. Inhibitory innervation of the walking legs of the scorpion Vaejovis spinigerus (Wood, 1863) (Vaejovidae, Scorpiones, Arachnida). Arthropod Struct. Dev. 31: 203-215
    • Harzsch S (2001) Neurogenesis in the crustacean ventral nerve cord: homology of neuronal stem cells in Malacostraca and Branchiopoda? Evol. Dev. 3: 154-169
    • Harzsch S, Walossek D (2001) Neurogenesis in the developing visual system of the branchiopod crustacean Triops longicaudatus (LeConte, 1846): corresponding patterns of compound-eye formation in Crustacea and Insecta? Dev. Genes Evol. 211: 37-43
    • Harzsch S, Waloszek D (2000) Serotonin-immunoreactive neurons in the ventral nerve cord of Crustacea: a character to study aspects of arthropod phylogeny. Arthropod Struct. Dev. 29: 307-322
    • Harzsch S, Benton J, Beltz B (2000) An unusual case of a mutant lobster embryo with double brain and double ventral nerve cord. Arthropod Struct. Dev. 29: 95-99
    • Harzsch S (2004) The arthropod tritocerebrum: a “non-drosophilocentric” perspective. Evol. Dev. 6: 303-309
    • Harzsch S (2003) Ontogeny of the ventral nerve cord in malacostracan crustaceans: a common plan for neuronal development in Crustacea, Hexapoda and other Arthropoda? Arthropod Struct. Dev. 32: 17-37.
    • Harzsch S (2003) Introduction - Development of the arthropod nervous system: variations on a common theme? Arthropod Struct. Dev. 32: 3-4. 48*
    • Harzsch S (2002) Neurobiologie und Evolutionsforschung: „Neurophylogenie“ und die Stammesgeschichte der Euarthropoda. Neuroforum: 267-273
    • Harzsch S (2002) In the realm of the many limbs. German Research 1/2002: 25-27
    • Harzsch S (2002) From stem cell to structure: neurogenesis in the CNS of decapod crustaceans. In: The Crustacean Nervous System, K. Wiese ed., Springer Verlag: 417-432
    • Harzsch S (2001) Im Reich der vielen Füße. Forschung, Magazin der Deutschen Forschungsgemeinschaft 4/2001: 7-9
    • Harzsch S (2001) Entwicklung des Nervensystems der Crustacea: ein Beitrag zur Phylogenie der Arthropoda. Habilitationsschrift, Universität Bielefeld, Fakulät für Biologie
    1993 - 1999
    • Harzsch S, Miller J, Benton J, Beltz B (1999) From embryo to adult: persistent neurogenesis and apoptotic cell death shape the developing crustacean deutocerebrum. J. Neurosci. 19: 3472-3485
    • Harzsch S, Benton J, Dawirs RR, Beltz B (1999) A new look at embryonic development of the visual system in decapod crustaceans: neuropil formation, neurogenesis and apoptotic cell death. J. Neurobiol. 39: 294-306
    • Schmidt M, Harzsch S (1999) Comparative analysis of neurogenesis in the central olfactory pathway of adult decapod crustaceans by in vivo BrdU-labelling. Biol. Bull. 196: 127-136
    • Harzsch S, Miller J, Benton J, Dawirs R, Beltz B (1998) Neurogenesis in the thoracic neuromeres of two crustaceans with different styles of metamorphic development. J. Exp. Biol. 201: 2465-2479
    • Harzsch S, Anger K, Dawirs RR (1997) Immunocytochemical detection of acetylated *-tubulin and Drosophila synapsin in the embryonic crustacean nervous system. Int. J. Dev. Biol. 41: 477–484
    • Harzsch S, Dawirs RR (1996) Neurogenesis in the developing crab brain: postembryonic generation of neurons persists beyond metamorphosis. J. Neurobiol. 29: 384-398
    • Harzsch S, Dawirs RR (1996) Development of neurons exhibiting FMRFamide-related immunoreactivity in the central nervous system of spider crab larvae (Hyas araneus L., Decapoda, Majidae). J. Crust. Biol. 16: 10-19
    • Harzsch S, Dawirs RR (1995/96) Maturation of the compound eyes and eyestalk ganglia during larval development of the brachyuran crustaceans Hyas araneus L. (Decapoda, Majidae) and Carcinus maenas L. (Decapoda, Portunidae). Zoology 99: 189-204
    • Harzsch S, Dawirs RR (1995) A developmental study of serotonin-immunoreactive neurons in the larval central nervous system of the spider crab Hyas araneus (Decapoda, Brachyura). Invertebr. Neurosci. 1: 53-65
    • Harzsch S, Dawirs RR (1994) Neurogenesis in larval stages of the spider crab Hyas araneus (Decapoda, Brachyura): proliferation of neuroblasts in the ventral nerve cord. Roux's Arch. Dev. Biol. 204: 93-100
    • Harzsch S, Dawirs RR (1993) On the morphology of the central nervous system in larval stages of Carcinus maenas L. (Decapoda, Brachyura). Helgoländer Meeresuntersuch. 47: 61-79
    • Harzsch S (1995) Zur Entwicklung des Zentralnervensystems der Krabben Carcinus maenas und Hyas araneus (Decapoda, Brachyura). Dissertation, Universität Bielefeld, Fakultät für Biologie
    • Harzsch S, Dawirs RR (1994) Development of the CNS in spider crab larvae. ZOEA Newsletter 1: 8-10

    Structure and Evolution of Invertebrate Nervous Systems

    edited by Andreas Schmidt-Rhaesa, Steffen Harzsch and Günter Purschke