Our publications on onychophorans

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


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.


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.