Our publications on onychophorans

(PDFs of non-open access articles available on request)


Martin, C., Jahn, H., Klein, M.M., Hammel, J.U., Stevenson, P.A., Homberg, U. & Mayer, G. (2022): The velvet worm brain unveils homologies and evolutionary novelties across panarthropods. BMC Biology 20:26. Free article.
Tait, N.N., Oliveira, I.S. & Mayer, G. (2022): Translation of N.B.K. Sänger’s (1871) “Peripatus capensis Gr. and Peripatus leuckartii n. sp.” (Onychophora), along with remarks about the author and significance of his work. Australian Zoologist 42:71–94. Free article.


Oliveira, I.S. & Mayer, G. (2021) Phylum Onychophora. In: B. Schierwater & R. DeSalle (Eds) Invertebrate Zoology: A Tree of Life Approach. CRC Press, Oxon, 459–475.


Martin, C., Hering, L., Metzendorf, N., Hormann, S., Kasten, S., Fuhrmann, S., Werckenthin, A., Herberg, F.W., Stengl, M. & Mayer, G. (2020): Analysis of pigment-dispersing factor neuropeptides and their receptor in a velvet worm. Frontiers in Endocrinology 11:273. Free article.


Oliveira, I.S.*, Kumerics, A.*, Jahn, H., Müller, M., Pfeiffer, F. & Mayer, G. (2019): Functional morphology of a lobopod: Case study of an onychophoran leg. Royal Society Open Science 6:191200. *Shared first authorship. Free article.
Baer, A., Oliveira, I.S., Schmidt, S., Harrington, M.J. & Mayer, G. (in press): Mechanismen der Natur als Vorbild für nachhaltige Biomaterialien: Einblicke in den „Sekundenkleber“ der Stummelfüßer. Biologie in unserer Zeit.
Baer, A., Schmidt, S., Mayer, G. & Harrington, M.J. (in press): Fibers on the fly: Multiscale mechanisms of fiber formation in the capture slime of velvet worms. Integrative and Comparative Biology.
Treffkorn, S., Lagos, O.Y.H. & Mayer, G. (2019): Evidence for cell turnover as the mechanism responsible for the transport of embryos towards the vagina in viviparous onychophorans (velvet worms). Frontiers in Zoology 16:16. Free article.
Baer, A., Horbelt, N., Nijemeisland, M., Garcia, S.J., Fratzl, P., Schmidt, S., Mayer, G. & Harrington, M.J. (2019): Shear-induced β-crystallite unfolding in condensed phase nanodroplets promotes fiber formation in a biological adhesive. ACS Nano 13:4992–5001.
Treffkorn, S. & Mayer, G. (2019): Expression of NK genes that are not part of the NK cluster in the onychophoran Euperipatoides rowelli (Peripatopsidae). BMC Developmental Biology 19:7. Free article.
McCravy, K.W. & Oliveira, I.S. (2019): Onychophorans (Velvet Worms): Natural history and conservation of a vulnerable invertebrate taxon in the Caribbean islands. Caribbean Naturalist Special Issue 2:135–155.
Segura, B. & Mayer, G. (2019): Onicóforos de la Cordillera de la Costa de Chile. In: C. Smith-Ramírez & F.A. Squeo (Eds) Biodiversidad y Ecología de los Bosques Costeros de Chile. Editorial Universidad de Los Lagos, Osorno, pp. 53–56.


Baer, A., Hänsch, S., Mayer, G., Harrington, M.J. & Schmidt, S. (2018): Reversible supramolecular assembly of velvet worm adhesive fibers via electrostatic interactions of charged phosphoproteins. Biomacromolecules 19: 4034–4043.
Ou, Q. & Mayer, G. (2018): A Cambrian unarmored lobopodian, †Lenisambulatrix humboldti gen. et sp. nov., compared with new material of †Diania cactiformis. Scientific Reports 8: 13667.
Schumann, I.*, Kenny, N., Hui, J., Hering, L.* & Mayer, G. (2018): Halloween genes in panarthropods and the evolution of the early moulting pathway in Ecdysozoa. Royal Society Open Science 5: 180888. *Equal contribution. Free article.
Oliveira, I.S., Ruhberg, H., Rowell, D.M. & Mayer, G. (2018): Revision of Tasmanian viviparous velvet worms (Onychophora: Peripatopsidae) with descriptions of two new species. Invertebrate Systematics 32: 909–932.
Treffkorn, S., Kahnke, L., Hering, L. & Mayer, G. (2018): Expression of NK cluster genes in the onychophoran Euperipatoides rowelli: Implications for the evolution of NK family genes in nephrozoans. EvoDevo 9: 17. Free article.
Kirwan, J.D., Graf, J., Smolka, J., Mayer, G., Henze, M.J. & Nilsson, D.-E. (2018): Low resolution vision in a velvet worm (Onychophora). Journal of Experimental Biology 221: jeb175802.
Jahn, H., Oliveira, I.S., Gross, V., Martin, C., Hipp, A., Mayer, G. & Hammel, J.U. (2018): Evaluation of contrasting techniques for X-ray imaging of velvet worms (Onychophora). Journal of Microscopy 270: 343–358.


Müller, M., Oliveira, I.S., Allner, S., Ferstl, S., Bidola, P., Mechlem, K., Fehringer, A., Hehn, L., Dierolf, M., Achterhold, K., Gleich, B., Hammel, J.U., Jahn, H., Mayer, G. & Pfeiffer, F. (2017): Myoanatomy of the velvet worm leg revealed by laboratory-based nanofocus X-ray source tomography. Proceedings of the National Academy of Sciences of the United States of America 114: 12378–12383.

Baer, A., Schmidt, S., Haensch, S., Eder, M., Mayer, G. & Harrington, M. (2017): Mechanoresponsive lipid-protein nanoglobules facilitate reversible fibre formation in velvet worm slime. Nature Communications 8: 974.
Martin, C., Gross, V., Hering, L., Tepper, B., Jahn, H., Oliveira, I.S., Stevenson, P.A. & Mayer, G. (2017): The nervous and visual systems of onychophorans and tardigrades: learning about arthropod evolution from their closest relatives. Journal of Comparative Physiology A 203: 565–590.
Oliveira, I.S. & Mayer, G. (2017): A new egg-laying onychophoran (Peripatopsidae) reveals evolutionary and biogeographical aspects of Australian velvet worms. Organisms Diversity & Evolution 17: 375–391.
Martin, C., Gross, V., Pflüger, H.J., Stevenson, P.A. & Mayer, G. (2017): Assessing segmental versus non-segmental features in the ventral nervous system of onychophorans (velvet worms). BMC Evolutionary Biology 17:3. Free article.
Treffkorn, S. & Mayer, G. (2017): Conserved versus derived patterns of controlled cell death during the embryonic development of two species of Onychophora (velvet worms). Developmental Dynamics 246:  403–416.
von Byern, J., Müller, C., Voigtländer, K., Dorrer, V., Marchetti-Deschmann, M., Flammang, P. & Mayer, G. (2017): Examples of Bioadhesives for Defence and Predation. In: S.N. Gorb & E.V. Gorb (Eds) Biologically-Inspired Systems 10: Functional Surfaces in Biology III: Diversity of the Physical Phenomena. Springer, Cham, pp. 141–191.


Oliveira, I.S.*, Bai, M.*, Jahn, H., Gross, V., Martin, C., Hammel, J.U., Zhang, W. & Mayer, G. (2016): Earliest onychophoran in amber reveals Gondwanan migration patterns. Current Biology 26: 2594–2601. *Shared first authorship.
Schumann, I., Hering, L. & Mayer, G. (2016): Immunolocalization of arthropsin in the onychophoran Euperipatoides rowelli (Peripatopsidae). Frontiers in Neuroanatomy 10:80. Free article
Mayer, G. (2016): Onychophora. In: A. Schmidt-Rhaesa, S. Harzsch & G. Purschke (Eds) Structure and Evolution of Invertebrate Nervous Systems. Oxford University Press, Oxford, pp. 390–401.


Martin, C. & Mayer, G. (2015): Insights into the segmental identity of post-oral commissures and pharyngeal nerves in Onychophora based on retrograde fills. BMC Neuroscience 16: 53. Free article.
Franke, F.A. & Mayer, G. (2015): Expression study of the hunchback ortholog in embryos of the onychophoran Euperipatoides rowelli. Development Genes and Evolution 225: 207–219.
Mayer, G., Hering, L., Stosch, J.M., Stevenson, P.A. & Dircksen, H. (2015): Evolution of pigment-dispersing factor neuropeptides in Panarthropoda: Insights from Onychophora (velvet worms) and Tardigrada (water bears). Journal of Comparative Neurology 523: 1865–1885
Mayer, G., Oliveira, I.S., Baer, A., Hammel, J.U., Gallant, J. & Hochberg, R. (2015): Capture of prey, feeding, and functional anatomy of the jaws in velvet worms (Onychophora). Integrative and Comparative Biology 55: 217–227.
Beckmann, H., Hering, L., Henze, M.J., Kelber, A., Stevenson, P.A. & Mayer, G. (2015): Spectral sensitivity in Onychophora (velvet worms) revealed by electroretinograms, phototactic behaviour and opsin gene expression. Journal of Experimental Biology 218: 915-922.
Oliveira, I.S., Lacorte, G.A., Weck-Heimann, A., Cordeiro, L.M., Wieloch, A.H. & Mayer, G. (2015): A new and critically endangered species and genus of Onychophora (Peripatidae) from the Brazilian savannah – a vulnerable biodiversity hotspot. Systematics and Biodiversity 13: 211–233.

Franke, F.A., Schumann, I., Hering, L. & Mayer, G. (2015): Phylogenetic analysis and expression patterns of Pax genes in the onychophoran Euperipatoides rowelli reveal a novel bilaterian Pax subfamily. Evolution & Development 17: 3–20.
Mayer, G., Franke, F.A., Treffkorn, S., Gross, V. & Oliveira, I.S. (2015): Onychophora. In: A. Wanninger (Ed.) Evolutionary Developmental Biology of Invertebrates 3: Ecdysozoa I: Non-tetraconata. Springer, Berlin, 53–98.


Mayer, G., Martin, C., Oliveira, I.S., Franke, F.A. & Gross, V. (2014): Latest anomalocaridid affinities challenged. Nature 516: E1–E2.
Franke, F.A. & Mayer, G. (2014): Controversies surrounding segments and parasegments in Onychophora: Insights from the expression patterns of four "segment polarity genes" in the peripatopsid Euperipatoides rowelli. PLOS ONE 9(12): e114383. Free article.
Murdock, D.J.E., Gabbott, S.E., Mayer, G. & Purnell, M.A. (2014): Decay of velvet worms (Onychophora), and bias in the fossil record of lobopodians. BMC Evolutionary Biology 14: 222. Free article.
Oliveira, I.S., Lüter, C., Wolf, K.W. & Mayer, G. (2014): Evolutionary changes in the integument of the onychophoran Plicatoperipatus jamaicensis (Peripatidae). Invertebrate Biology 133: 274–280.
Baer, A., Oliveira, I.S., Steinhagen, M., Beck-Sickinger, A.G. & Mayer, G. (2014): Slime protein profiling: A non-invasive tool for species identification in Onychophora (velvet worms). Journal of Zoological Systematics and Evolutionary Research 52: 265–272.
Martin, C. & Mayer, G. (2014): Neuronal tracing of oral nerves in a velvet worm – Implications for the evolution of the ecdysozoan brain. Frontiers in Neuroanatomy 8: 7. Free article.
Murienne, J., Daniels, S.R., Buckley, T.R., Mayer, G. & Giribet, G. (2014): A living fossil tale of Pangean biogeography. Proceedings of the Royal Society B: Biological Sciences 281: 1471–2954.


Oliveira, I.S., Tait, N.N., Strübing, I. & Mayer, G. (2013): The role of ventral and preventral organs as attachment sites for segmental limb muscles in Onychophora. Frontiers in Zoology 10: 73. Free article.
Mayer, G., Martin, C., Rüdiger, J., Kauschke, S., Stevenson, P.A., Poprawa, I., Hohberg, K., Schill, R.O., Pflüger, H.-J. & Schlegel, M. (2013): Selective neuronal staining in tardigrades and onychophorans provides insights into the evolution of segmental ganglia in panarthropods. BMC Evolutionary Biology 13: 230. Free article.

Mayer, G., Kauschke, S., Rüdiger, J. & Stevenson, P.A. (2013): Neural markers reveal a one-segmented head in tardigrades (water bears). PLOS ONE 8(3): e59090. Free article.
Oliveira, I.S. & Mayer, G. (2013): Apodemes associated with limbs support serial homology of claws and jaws in Onychophora (velvet worms). Journal of Morphology 274: 1180–1190.
Treffkorn, S. & Mayer, G. (2013): Expression of the decapentaplegic ortholog in embryos of the onychophoran Euperipatoides rowelli. Gene Expression Patterns 13: 384–394. Free article.
Oliveira, I.S., Schaffer, S., Kvartalnov, P.V., Galoyan, E.A., Palko, I.V., Weck-Heimann, A., Geißler, P., Ruhberg, H. & Mayer, G. (2013): A new species of Eoperipatus (Onychophora) from Vietnam reveals novel morphological characters for the South-East Asian Peripatidae. Zoologischer Anzeiger 252: 495–510.

Mayer, G. & Oliveira, I.S. (2013): Phylum Onychophora Grube, 1853. In: Zhang, Z.-Q. (Ed.) Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness (Addenda 2013). Zootaxa 3703: 15–16. Free article.
Ruhberg, H. & Mayer, G. (2013): Onychophora, Stummelfüßer. In: W. Westheide & G. Rieger (Eds) Spezielle Zoologie. Teil 1. Einzeller und Wirbellose Tiere. Springer-Verlag, Berlin, pp. 457–464.
Stein, M. & Mayer, G. (2013): Fossile Panarthropoda. In: W. Westheide & G. Rieger (Eds) Spezielle Zoologie. Teil 1. Einzeller und Wirbellose Tiere. Springer-Verlag, Berlin, pp. 472–473.


Jeffery, N.W., Oliveira, I.S., Gregory, T.R., Rowell, D.M. & Mayer, G. (2012): Genome size and chromosome number in velvet worms (Onychophora). Genetica 140: 497–504.
Oliveira, I.S., Franke, F.A., Hering, L., Schaffer, S., Rowell, D.M., Weck-Heimann, A., Monge-Nájera, J., Morera-Brenes, B. & Mayer, G. (2012): Unexplored character diversity in Onychophora (velvet worms): a comparative study of three peripatid species. PLOS ONE 7(12): e51220. Free article.

Ou, Q., Shu, D. & Mayer, G. (2012): Cambrian lobopodians and extant onychophorans provide new insights into early cephalization in Panarthropoda. Nature Communications 3: 1261. Free article.
Haug, J.T., Mayer, G., Haug, C. & Briggs, D.E.G. (2012): A Carboniferous non-onychophoran lobopodian reveals long-term survival of a Cambrian morphotype. Current Biology 22: 1673–1675. Free article.
Hering, L., Henze, M.J., Kohler, M., Kelber, A., Bleidorn, C., Leschke, M., Nickel, B., Meyer, M., Kircher, M., Sunnucks, P. & Mayer, G. (2012): Opsins in Onychophora (velvet worms) suggest a single origin and subsequent diversification of visual pigments in arthropods. Molecular Biology and Evolution 29: 3451–3458. Free article.

Baer, A. & Mayer, G. (2012): Comparative anatomy of slime glands in Onychophora (velvet worms). Journal of Morphology 273: 1079–1088.
Oliveira, I.S., Read, V.M.St.J. & Mayer, G. (2012): A world checklist of Onychophora (velvet worms), with notes on nomenclature and status of names. ZooKeys 211: 1–70. Free article.


Oliveira, I.S., Lacorte, G.A., Fonseca, C.G., Wieloch, A.H. & Mayer, G. (2011): Cryptic speciation in Brazilian Epiperipatus (Onychophora: Peripatidae) reveals an underestimated diversity among the peripatid velvet worms. PLOS ONE 6(6): e19973. Free article.
Mayer, G. & Oliveira, I. S. (2011): Phylum Onychophora Grube, 1853. In: Zhang, Z.-Q. (Ed.) Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness. Zootaxa 3148: 98. Free article.
Lacorte, G.A., Oliveira, I.S. & Fonseca, C.G. (2011): Population structure and demographic inferences of the endangered onychophoran species Epiperipatus acacioi (Onychophora: Peripatidae). Genetics and Molecular Research 10: 2775–2785. Free article.
Whitington, P.M. & Mayer, G. (2011): The origins of the arthropod nervous system: insights from the Onychophora. Arthropod Structure & Development 40: 193–209. Invited paper for a Special Issue.
Chisholm, R.H., Hughes, B.D., Landman, K.A., Mayer, G. & Whitington, P.M. (2011): When are cellular oscillators sufficient for sequential segmentation? Journal of Theoretical Biology 279: 150–160.
Lacorte, G.A., Oliveira, I.S. & Fonseca, C.G. (2011): Phylogenetic relationships among the Epiperipatus lineages (Onychophora: Peripatidae) from the Minas Gerais State, Brazil. Zootaxa 2755: 57–65.
Groh, C. & Mayer, G. (2011): Evolution of the arthropod nervous system. Part 1. Arthropod Structure & Development 40: 191–192. Editorial for a Special Issue.
Groh, C. & Mayer, G. (2011): Evolution of the arthropod nervous system. Part 2. Arthropod Structure & Development 40: 303. Editorial for a Special Issue.


Allwood, J., Gleeson, D., Mayer, G., Daniels, S., Beggs, J., & Buckley, T.R. (2010): Support for vicariant origins of the New Zealand Onychophora fauna. Journal of Biogeography 37: 669–681.
Mayer, G., Whitington, P.M., Sunnucks, P. & Pflüger, H.-J. (2010): A revision of brain composition in Onychophora (velvet worms) suggests that the tritocerebrum evolved in arthropods. BMC Evolutionary Biology 10: 255. Free article.
Mayer, G., Kato, C., Quast, B., Chisholm, R.H., Landman, K.A. & Quinn, L.M. (2010): Growth patterns in Onychophora (velvet worms): lack of a localised posterior proliferation zone. BMC Evolutionary Biology 10: 339. Free article.
*Braband, A., *Cameron, S.L., Podsiadlowski, L., Daniels, S.R. & Mayer, G. (2010): The mitochondrial genome of the onychophoran Opisthopatus cinctipes (Peripatopsidae) reflects the ancestral mitochondrial gene arrangement of Panarthropoda and Ecdysozoa. *Equal contribution. Molecular Phylogenetics & Evolution 57: 285–292.

Braband, A., Podsiadlowski, L., Cameron, S.L., Daniels, S.R. & Mayer, G. (2010): Extensive duplication events account for multiple control regions and pseudogenes in the mitochondrial genome of the velvet worm Metaperipatus inae (Onychophora, Peripatopsidae). Molecular Phylogenetics & Evolution 57: 293–300.

Oliveira, I.S., Wieloch, A.H. & Mayer, G. (2010): Revised taxonomy and redescription of two species of the Peripatidae (Onychophora) from Brazil: a step towards consistent terminology of morphological characters. Zootaxa 2493: 16–34.


Mayer, G. & Whitington, P.M. (2009): Velvet worm development links myriapods with chelicerates. Proceedings of the Royal Society B: Biological Sciences 276: 3571–3579. Free article. Recommended on Faculty of 1000.
Mayer, G. & Whitington, P.M. (2009): Neural development in Onychophora (velvet worms) suggests a step-wise evolution of segmentation in the nervous system of Panarthropoda. Developmental Biology 335: 263–275. Free article.
Mayer, G. & Tait, N.N. (2009): Position and development of oocytes in velvet worms shed light on the evolution of the ovary in Onychophora and Arthropoda. Zoological Journal of the Linnean Society 157: 17–33.
Podsiadlowski, L., Braband, A. & Mayer, G. (2008): The complete mitochondrial genome of the onychophoran Epiperipatus biolleyi reveals a unique transfer RNA set and provides further support for the Ecdysozoa hypothesis. Molecular Biology and Evolution 25: 42–51. Free article.

Mayer, G. & Harzsch, S. (2008): Distribution of serotonin in the trunk of Metaperipatus blainvillei (Onychophora, Peripatopsidae): Implications for the evolution of the nervous system in Arthropoda. Journal of Comparative Neurology 507: 1196–1208.
Oliveira, I.S. & Wieloch, A.H. (2008): Testemunhas da evolução. Ciência Hoje 41: 66–68. Free article.
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. Free article.
Maas, A., Mayer, G., Kristensen, R.M., & Waloszek, D. (2007): A Cambrian micro-lobopodian and the evolution of arthropod locomotion and reproduction. Chinese Science Bulletin 52: 3385–3392.
Mayer, G. (2007): Metaperipatus inae sp. nov. (Onychophora: Peripatopsidae) from Chile with a novel ovarian type and dermal insemination. Zootaxa 1440: 21–37.
Mayer, G. (2006): Structure and development of the onychophoran eyes – what is the ancestral visual organ in arthropods? Arthropod Structure & Development 35: 231–245. Invited paper for a Special Issue.
Mayer, G. (2006): Origin and differentiation of nephridia in the Onychophora provide no support for the Articulata. Zoomorphology 125: 1–12.
Oliveira, I.S. & Wieloch, A.H. (2005): Macroperipatus machadoi sp. n. (Onychophora: Peripatidae) da Floresta Atlântica de Minas Gerais, Brasil. Lundiana 6: 61–66. Free article.
Mayer, G. & Koch, M. (2005): Ultrastructure and fate of the nephridial anlagen in the antennal segment of Epiperipatus biolleyi (Onychophora, Peripatidae) – Evidence for onychophoran antennae being modified legs. Arthropod Structure & Development 34: 471–480.
Mayer, G., Bartolomaeus, T. & Ruhberg, H. (2005): Ultrastructure of mesoderm in embryos of Opisthopatus roseus (Onychophora, Peripatopsidae): Revision of the "Long Germ Band" hypothesis for Opisthopatus. Journal of Morphology 263: 60–70.
Mayer, G., Ruhberg, H. & Bartolomaeus, T. (2004): When an epithelium ceases to exist – An ultrastructural study on the fate of the embryonic coelom in Epiperipatus biolleyi (Onychophora, Peripatidae). Acta Zoologica 85: 163–170.