Onychophorology, the study of velvet worms, historical trends, landmarks, and researchers from 1826 to 2020 (a literature review)





history of science, the study of invertebrates, research patterns, studies on onychophorans


Velvet worms, also known as peripatus or onychophorans, are a phylum of evolutionary importance that has survived all mass extinctions since the Cambrian period. They capture prey with an adhesive net that is formed in a fraction of a second. The first naturalist to formally describe them was Lansdown Guilding (1797-1831), a British priest from the Caribbean island of Saint Vincent. His life is as little known as the history of the field he initiated, Onychophorology. This is the first general history of Onychophorology, which has been divided into half-century periods. The beginning, 1826-1879, was characterized by studies from former students of famous naturalists like Cuvier and von Baer. This generation included Milne-Edwards and Blanchard, and studies were done mostly in France, Britain, and Germany. In the 1880-1929 period, research was concentrated on anatomy, behavior, biogeography, and ecology; and it is in this period when Bouvier published his mammoth monograph. The next half-century, 1930-1979, was important for the discovery of Cambrian species; Vachon’s explanation of how ancient distribution defined the existence of two families; DNA and electron microscopy from Brazil; and primitive attempts at systematics using embryology or isolated anatomical characteristics. Finally, the 1980-2020 period, with research centered in Australia, Brazil, Costa Rica, and Germany, is marked by an evolutionary approach: from body and behavior to geographic distribution; the discovery of how they form their adhesive net; the reconstruction of Cambrian onychophoran communities, the first experimental taphonomy; the first country-wide map of conservation status (in Costa Rica); the first model of why they survive in cities; the discovery of new phenomena like food hiding, parental feeding investment, and ontogenetic diet shift; and the birth of a new research branch, onychophoran ethnobiology. While a few names often appear in the literature, most knowledge was produced by a mass of researchers who entered the field only briefly.


[Anonymous]. (1885). Peripatus. In Report on the Scientific Results of the Voyage of H.M.S. Challenger During the Years 1873-76 (pp. 284-286). London: Longmans & Co.

[Anonymous]. (1895). Report of club meetings, 19 April 1895. Journal of the Trinidad Field Naturalists' Club, 2, 187-189.

[Anonymous]. (2012). Eugène Louis Bouvier (1856-1944). Paris: Institut Pasteur. Retrieved from https://webext.pasteur.fr/archives/buv0.html

Alexander, A. (1958). Peripatus: Fierce little giant. Animal Kingdom, 61, 122-125

Altincicek, B., & Vilcinskas, A. (2008). Identification of immune inducible genes from the velvet worm Epiperipatus biolleyi (Onychophora). Developmental and Comparative Immunology, 32(12), 1416-21. https://doi.org/10.1016/j.dci.2008.06.004

Annandale, N. (1912). The occurrence of Peripatus on the North-East frontier of India. Nature, 88, 449. https://doi.org/10.1038/088449b0

Arnett, R. H. (1947). Epiperipatus braziliensis (Bouvier) on Barro Colorado Island, Canal Zone. Entomological News, 58, 59-60.

Arvy, L. (1954). Présentation de documents sur la leucopoièse chez Peripatopsis capensis Grube. Bulletin de la Société Zoologique de France, 79, 13.

Audouin, M. & Milne-Edwards, H. (1833). Classification des Annélides, et description de celles qui habitent les cites de la France - sixième Famille: Péripatiens. Annales des Sciences Naturelles, 30, 411-414. https://doi.org/10.5962/bhl.part.8010

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(10), 4034-4043. DOI: 10.1021/acs.biomac.8b01017

Bai, T. T., & Anh, N. D. (2012). Discovery of Eoperipatus sp. (Peripatidae), the first representative of Onychophora in Vietnam. TAP CHI SINH HOC, 32(4), 36-39. DOI: 10.15625/0866-7160/v32n4.718

Balfour, F.M. (1883). The anatomy and development of Peripatus capensis. Quarterly Journal of Microscopical Science, 23, 213-259.

Ballard, J. W. O.; Olsen, G. J.; Faith, D. P., Odgers; W. A., Rowell, D. M., & Atkinson, P. W. (1992). Evidence from 12S ribosomal RNA sequences that onychophorans are modified arthropods. Science, 258, 1345-1348. https://doi.org/10.1126/science.1455227

Barquero-González, J. P.; Morera-Brenes, B., & Monge-Nájera, J. (2018). The relationship between humidity, light and the activity pattern of a velvet worm, Epiperipatus sp. (Onychophora: Peripatidae), from Bahía Drake, South Pacific of Costa Rica. Brazilian Journal of Biology, 78(3), 408-413. https://doi.org/10.1590/1519-6984.166495

Barquero-González, J. P.; Vega-Hidalgo, A., & Monge-Nájera, J. (2019). Feeding behavior of Costa Rican velvet worms: food hiding, parental feeding investment and ontogenetic diet shift (Onychophora: Peripatidae). UNED Research Journal, 11(2), 85-88. DOI: 10.22458/urj.v11i2.2195

Barrett, D. (2013). Multiple-scale resource selection of an undescribed urban invertebrate (Onychophora: Peripatopsidae) in Dunedin, New Zealand (Master´s Thesis). New Zealand, University of Otago. Retrieved from https://ourarchive.otago.ac.nz/handle/10523/4044

Bell, F. J. (1887). Habitat of Peripatus leuckarti. Annals and Magazine of Natural History [Series 5], 20, 252. https://doi.org/10.1080/00222938709460049

Bellomy, M. D. (1955). Peripatus - between worm and insect. Frontiers (Philadelphia, Pennsylvania, USA), 20, 42-44.

Benkendorff, K.; Beardmore, K.; Gooley, A. A.; Packer, N. H., & Tait, N. N. (1999). Characterisation of the slime gland secretion from the peripatus, Euperipatoides kanangrensis (Onychophora: Peripatopsidae). Comparative Biochemistry and Physiology, Part B, 124, 457-465. https://doi.org/10.1016/S0305-0491(99)00145-5

Birket-Smith, S. J. R. (1974). The anatomy of the body wall of Onychophora. Zoologische Jahrbücher, Abteilung für Anatomie und Ontogenie der Tiere, 93, 123-154.

Blanchard, E. (1847). Recherches sur l'organisation des Vers. Annales des Sciences Naturelles [3e Série], 8, 119-149. https://doi.org/10.5962/bhl.title.51506

Blanchard, E. (1849). Malacopodes. In C. Gay (Ed.), Historia Fisica y Politica de Chile (Vol. 23): Zoologia, Tomo Tercero (pp. 57-60). Paris: privately published.

Blanckaert, C. & Hurel, A. (2017). Louis-Eugène Bouvier (1856-1944), un amateur à la chaire d’entomologie? Retrieved from https://objethistoire.hypotheses.org/953

Boas, I. E. V. (1898). Om Peripatus stilling i dyreriget. Kongelige Danske Videnskabernes Selskabs Forhandlinger, 6, 345-365.

Bouvier, E. L. (1902). Observations nouvelles sur l'évolution et l'origine des Péripates. Comptes Rendus Hebdomadaires des Seances de l'Academie des Sciences, 134, 55-58. https://doi.org/10.5962/bhl.part.19865

Bouvier, E. L. (1905). Monographie des Onychophores. Annales des Sciences Naturelles, Zoologie et Biologie Animale [9e Série], 2, 1-383.

Bouvier, E. L. (1907). Monographie des Onychophores. Annales des Sciences Naturelles, Zoologie et Biologie Animale [9e Série], 5, 61-318

Bouvier, E. L. (1928). A propos des observations du Fr. Claude-Joseph sur un Péripate du Chili. Annales des Sciences Naturelles, Zoologie et Biologie Animale [10e Série], 11, 260.

Briscoe, D. A., & Tait, N. N. (1995). Allozyme evidence for extensive and ancient radiations in Australian Onychophora. Zoological Journal of the Linnean Society, 114, 91-102. https://doi.org/10.1111/j.1096-3642.1995.tb00114.x

Brues, C. T. (1935). Varietal forms of Peripatus in Haiti. Psyche, 42, 58-62. https://doi.org/10.1155/1935/56795

Brues, C. T. (1941). Peripatus (Macroperipatus) geayi in Panama. Psyche, 48, 111-112. https://doi.org/10.1155/1941/34357

Bruntz, L. (1903). Excrétion et phagocytose chez les Onychophores. Comptes Rendus Hebdomadaires des Seances de l'Academie des Sciences, 136, 1148-1150. https://doi.org/10.5962/bhl.part.3483

Camerano, L. (1896). Onicofori raccolti nel Darien da Dott. E. Festa. Bollettino del Museo Regionale di Scienze Naturali Torino, 11, 1-2

Campiglia, S. S. (1969). Tegumento, muda e ciclo de intermuda em Peripatus acacioi Marcus e Marcus (onicoforo) (Dissertation thesis). Universidade de Sao Paulo, Brazil.

Carus, J. V. (1863). Onychophora. In J. V. Carus & A. Gerstaecker (Eds.), Handbuch der Zoologie (pp. 446-447). Leipzig, Germany: Verlag von Wilhelm Engelmann.

Clark, A. H. & Zetek, J. (1946). The onychophores of Panama and the Canal Zone. Proceedings of the United States National Museum, 96, 205-213. https://doi.org/10.5479/si.00963801.96-3197.205

Clark, A. H. (1915). The present distribution of the Onychophora, a group of terrestrial invertebrates. Smithsonian Miscellaneous Collections, 65, 1-25.

Concha, A., Mellado, P., Morera-Brenes, B., Sampaio Costa, C., Mahadevan, L., & Monge-Nájera, J. (2015). Oscillation of the velvet worm slime jet by passive hydrodynamic instability. Nature Communications, 6, 6292. https://doi.org/10.1038/ncomms7292

Corrales-Urena, Y. R., Sanchez, A., Pereira, R., Rischka, K., Kowalik, T., & Vega-Baudrit, J. (2017). Extracellular micro and nanostructures forming the velvet worm solidified adhesive secretion. Materials Research Express, 4(12). https://doi.org/10.1088/2053-1591/aa9940

Costa, C. S. (2016). Sistemática e análise filogenética de Epiperipatus Clark, 1913 baseada em dados moleculares e morfológicos (Onychophora: Peripatidae) (Tese de Doutorado). Universidade de São Paulo, São Paulo. DOI: 10.11606/T.41.2016.tde-25082016-100135

Costa, C. S., & Giribet, G. (2016). Taxonomic Notes on Mesoperipatus tholloni (Onychophora: Peripatidae), an Elusive Velvet Worm from Gabon. Breviora, 552(1), 1-10. DOI: 10.3099/MCZ30.1

Costa, C. S.; Chagas-Junior, A.; Pinto-da-Rocha, R.; Costa, C. S.; Chagas-Junior, A., & Pinto-da-Rocha, R. (2018). Redescription of Epiperipatus edwardsii, and descriptions of five new species of Epiperipatus from Brazil (Onychophora: Peripatidae). Zoologia, 35, 1-15. DOI: 10.3897/zoologia.35.e23366

Craig, C. L. (1997). Evolution of arthropod silks. Annual Review of Entomology, 42, 231-267. https://doi.org/10.1146/annurev.ento.42.1.231

Crampton, G. C. (1928). The evolution of the head region in lower arthropods and its bearing upon the origin and relationships of the arthropodan groups. Canadian Entomologist, 60, 284-301. https://doi.org/10.4039/Ent60284-12

Cunha, W. T. R.; Santos, R. C. O.; Araripe, J.; Sampaio, I.; Schneider, H.; & Rêgo, P. S. (2017). Molecular analyses reveal the occurrence of three new sympatric lineages of velvet worms (Onychophora: Peripatidae) in the eastern Amazon basin. Genetics and Molecular Biology, 40(1), 147-152. DOI: 10.1590/1678-4685-gmb-2016-0037

Cupul-Magaña, F. G., & Navarrete-Heredia, J. (2008). Rediscovery and New Data for Oroperipatus eisenii (Wheeler, 1898) from Mexico (Onychophora: Peripatidae). Entomological News, 119(5), 545-549. DOI: 10.3157/0013-872x-119.5.545

Curach, N., & Sunnucks, P. (1999). Molecular anatomy of an onychophoran: Compartmentalized sperm storage and heterogeneous paternity. Molecular Ecology, 8, 1375-1385. https://doi.org/10.1046/j.1365-294x.1999.00698.x

Dakin, W. J. (1921). The eye of Peripatus. Quarterly Journal of Microscopical. Science, 65, 163-172.

Dakin, W. J. (1922). The infra-cerebral organs of Peripatus. Quarterly Journal of Microscopical Science, 66, 409-417.

Dakin, W. J., & Fordham, M. G. C. (1926). Birth of Peripatus in England. Nature, 117, 858. https://doi.org/10.1038/117858b0

Darteville, E. (1958). Les Péripates. Congo-Tervuren, 2, 65-68.

De Haro, A. (1998). Origen y relaciones fitogenéticas entre Artrópodos, Onicóforos, Anélidos y Lofoforados, según datos moleculares y morfológicos. Boletín de la Real Sociedad Española de Historia Natural Sección Biológica, 94, 103-113.

De la Fuente, J. A. (1975). Esquema filogenético de la línea Onychophora-Myriapoda-Hexapoda. Boletín de la Real Sociedad Española de Historia Natural Sección Biológica, 73, 85-97.

De Quatrefages, A. (1848). Études sur les types inférieurs de l`embranchment des annelés. Mémoire sur la famille des Hermelliens (Hermellea Nob.). Annales des Sciences Naturelles [3e Série], 10, 5-58. https://doi.org/10.5962/bhl.part.26122

Delle Cave, L., & Simonetta, A. M. (1975). Notes on the morphology and taxonomic position of Aysheaia (Onychophora?) and of Skania (undetermined phylum). Monitore Zoologico Italiano [Nuéva Sèrie], 9, 67-81.

Dendy, A. (1889). Peripatus in Victoria. Victorian Naturalist. 5, 134-135. https://doi.org/10.1038/039366a0

Duboscq, O. (1920). Notes sur Opisthopatus cinctipes Purc. I. Sur les poils des papilles primaires et leur développement - II. Les organes ventraux du cerveau. Archives de Zoologie Experimentale et Generale,59, 21-27.

Duerden, J. E. (1901). Abundance of Peripatus in Jamaica. Nature, 63, 440-441. https://doi.org/10.1038/063440d0

Dunn, E. R. (1943). Zoological results of the Azuero Peninsula Panama Expedition of 1940. Part I - A new species of Peripatus. Notulae Naturae, 123, 1-5.

Dutra, D. D.; Cordeiro, L. M., & Araujo, D. (2018). Maior número cromossômico em Onychophora e indício de fusão cromossômica detectada por FISH telomérica. Semina: Ciências Biológicas e Da Saúde, 38(1supl), 199. DOI: 10.5433/1679-0367.2017V38N1SUPLP199

Eliott, S.; Tait, N. N., & Briscoe, D. A. (1993). A pheromonal function for the crural glands of the onychophoran Cephalofovea tomahmontis (Onychophora: Peripatopsidae). Journal of Zoology, 231, 1-9. https://doi.org/10.1111/j.1469-7998.1993.tb05348.x

Eriksson, B. J., & Budd, G. E. (2003). The cephalic nerves of the Onychophora and their bearing on our understanding of head segmentation and stem-group evolution of Arthropoda. Retrieved from http://www.diva-portal.org/smash/record.jsf?pid=diva2%3A68261&dswid=-803

Eriksson, B. J., Samadi, L., & Schmid, A. (2013). The expression pattern of the genes engrailed, pax6, otd and six3 with special respect to head and eye development in Euperipatoides kanangrensis Reid 1996 (Onychophora: Peripatopsidae). Development Genes and Evolution, 223(4), 237-246. DOI: 10.1007/s00427-013-0442-z

Espinasa, L.; Garvey, R.; Espinasa, J.; Fratto, C. A.; Taylor, S. J.; Toulkeridis, T., & Addison, A. (2015). Cave dwelling Onychophora from a lava tube in the Galapagos. Subterranean Biology, 15(1), 1-10. https://doi.org/10.3897/subtbiol.15.8468

Ewer, D.W., & van der Berg, R. (1954). A note on the pharmacology of the dorsal musculature of Peripatopsis. Journal of Experimental Biology, 31, 497-500.

Fedorov, B. G. (1927). On the morphology of the brain of Peripatus. In N. Sewertzoff & B. S. Matveiev (Eds.), Proceedings of the Second Congress of Zoologists, Anatomists, and Histologists of USSR: Moscow 4-10 May 1925 (pp. 92-94). Moscow: "Glavnauka".

Fletcher, J. J. (1891). Note on the supposed oviparity of P. leuckartii. Proceedings of the Linnean Society of New South Wales, 2nd Series, 6, 577.

Fletcher, J. J. (1895). On the specific identity of the Australian Peripatus, usually supposed to be P. leuckarti, Saenger. Proceedings of the Linnean Society of New South Wales, 2nd Series, 10, 172-194. https://doi.org/10.5962/bhl.part.24340

Gabe, M. (1956). Particularités histologiques des glandes crurales des Onychophores. Bulletin de la Société Zoologique de France, 81, 170.

Gaffron, E. (1883). Anatomie und Histologie von Peripatus (Dissertation thesis). Poland, University of Breslau.

Gardner, C. R.; Robson, E. A., & Stanford, C. (1978). The presence of monoamines in the nervous system of Peripatopsis (Onychophora). Experientia, 34, 1577-1578. https://doi.org/10.1007/BF02034683

Gatenby, J. B. (1959). A note on the spermiogenesis of Peripatoides novae-zealandiae. Transactions of the Royal Society of New Zealand, 87, 51-53.

Gegenbaur, C. (1874). Dritter Abschnitt. Würmer. In Grundriss der Vergleichenden Anatomie Verlag von Wilhelm Engelmann, Leipzig (pp. 122-204). https://doi.org/10.5962/bhl.title.2155

Gervais, P. (1836). Note déscriptive sur Peripatus brevis. Bulletin de la Société Entomologique de France [1836], 15.

Gleeson, D. (1996). Uncovering the radiation of New Zealand's Onychophora using molecular data. Proceedings of the International Congress of Entomology, 19, 61.

Goodrich, E. S. (1897). On the relation of the arthropod head to the annelid prostomium. Journal of Cell Science, 2(40), 247-268.

Gowri, N., & Sundara Rajulu, G. (1976). A comparative study of the organic components of the haemolymph of Eoperipatus weldoni (Onychophora) and a pill-millipede Arthrosphaera lutescens (Diplopoda). Journal of Animal Morphology and Physiology, 23, 60-75.

Grabham, M., & Cockerell, T. D. A. (1892). Peripatus re-discovered in Jamaica. Nature, 46, 514. https://doi.org/10.1038/046514a0

Graham, L. D., Glattauer, V., Li, D., Tyler, M. J., & Ramshaw, J. A. M. (2013). The adhesive skin exudate of Notaden bennetti frogs (Anura: Limnodynastidae) has similarities to the prey capture glue of Euperipatoides sp. velvet worms (Onychophora: Peripatopsidae). Comparative Biochemistry and Physiology. Part B, Biochemistry & Molecular Biology, 165(4), 250-9. DOI: 10.1016/j.cbpb.2013.04.008

Grenier, J. K.; Garber, T. L.; Warren, R.: Whitington, P. M., & Carroll, S. (1997). Evolution of the entire arthropod Hox gene set predated the origin and radiation of the onychophoran/arthropod clade. Current Biology, 7, 547-553. https://doi.org/10.1016/S0960-9822(06)00253-3

Grube, E. (1853). Über den Bau von Peripatus edwardsii. Müller's Archives of Anatomy and Physiology [1853], 322-360.

Guilding, L. (1826). Mollusca Caribbaeana. Zoological Journal, 2, 437-449.

Haase, E. (1889). Über die Bewegungen von Peripatus. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin [1889], 148-151. https://doi.org/10.5962/bhl.part.26115

Hackman, R. H., & Goldberg, M. (1975). Peripatus: its affinities and its cuticle. Science, 190, 582-583. https://doi.org/10.1126/science.190.4214.582

Harris, A. C. (1991). A large aggregation of Peripatoides novaezealandiae (Hutton, 1876) (Onychophora: Peripatopsidae). Journal of the Royal Society of New Zealand, 21, 405-406. https://doi.org/10.1080/03036758.1991.10420836

Havel, J. E.; Wilson, C. C., & Hebert, P. D. N. (1989). Parental investment and sex allocation in a viviparous onychophoran. Oikos, 56, 224-232. https://doi.org/10.2307/3565340

Hebert, P. D. N.; Billington, N.; Finston, T. L.; Boileau, M. G.; Beaton, M. J., & Barrette, R. J. (1991). Genetic variation in the onychophoran Plicatoperipatus jamaicensis. Heredity, 67, 221-229. https://doi.org/10.1038/hdy.1991.83

Heffron, J. J. A., Hepburn, H. R. & Zwi, L. J. (1977). Enzymic activities of the smooth body-wall muscle of the onychophoran Peripatopsis moseleyi. Biochemical Society Transactions, 5, 1748-1750. https://doi.org/10.1042/bst0051748

Heffron, J. J. A.; Hepburn, H. R., & Zwi, J. (1976). On the sarcoplasmic reticulum of onychophoran somatic smooth muscle. Naturwissenschaften, 63, 95. https://doi.org/10.1007/BF00622418

Hendrickson, J. (1957). Peripatus in Malaysia. Malayan Nature Journal, 12, 33-35.

Hepburn, H. R. & Heffron, J. J. A. (1976). On the skeletal collagen of an onychophoran, Peripatopsis mosleyi. Cytobiologie, 12, 481-486

Hill, R. P. (1950). Peripatus: A missing link. Discovery, 11, 14-18.

Hilton, W. A. (1946). Remarks on the habitat of Peripatus on Barro Colorado Island, C.Z. Journal of Entomology and Zoology [Claremont, California], 38, 27.

Horst, R. (1886). On a specimen of Peripatus, Guild., from Sumatra. Notes from the Leyden Museum, 8, 37-41.

Horst, R. (1910). Paraperipatus lorentzi Horst, a new Peripatus from Dutch New Guinea. Notes from the Leyden Museum, 32, 217-218.

Hou, X. G. & Bergström, J. (1991). The arthropods of the Lower Cambrian Chengjiang fauna, with relationships and evolutionary significance. In A. M. Simonetta & S. Conway Morris (Eds.), The Early Evolution of the Metazoa and the Significance of Problematic Taxa (pp. 179-187). Cambridge, England: Cambridge University Press.

Howard, R. A., & Howard, E. S. (1985). "The Reverend Lansdown Guilding, 1797-1831". Journal of the Arnold Arboretum, 37(4), 401-402.

Hoyle, G., & del Castillo, J. (1979). Neuromuscular transmission in Peripatus. Journal of Experimental Biology, 83, 13-29.

Hoyle, G., & Williams, M. (1980). The musculature of Peripatus and its innervation. Philosophical Transactions of the Royal Society B, Biological Sciences, 288, 481-510. https://doi.org/10.1098/rstb.1980.0024

Hutchinson, G. E. (1930). Restudy of some Burgess Shale fossils. Proceedings of the United States National Museum, 78, 1-25. https://doi.org/10.5479/si.00963801.78-2854.1

Hutchinson, G. E. (1969). Aysheaia and the general morphology of the Onychophora. American Journal of Science, 267, 1062-1066. https://doi.org/10.2475/ajs.267.9.1062

Janssen, R., & Budd, G. E. (2017). Investigation of endoderm marker-genes during gastrulation and gut-development in the velvet worm Euperipatoides kanangrensis. Developmental Biology, 427(1), 155-164. DOI: 10.1016/j.ydbio.2017.04.014

Janvier, H. (1928). Le régime de l'Opisthopatus blainvillei. Comptes Rendus Hebdomadaires des Seances de l'Academie des Sciences, 186, 1748-1749.

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(10-12), 497-504. DOI: 10.1007/s10709-013-9698-5

Jerez-Jaimes, J. H., & Bernal-Pérez, M. C. (2009). Taxonomía de onicóforos de Santander, Colombia y termogravimetría, calorimetría de barrido diferencial y espectroscopía infrarroja de la secreción adhesiva (Onychophora: Peripatidae). Revista de Biologia Tropical, 57(3), 567-588. https://doi.org/10.15517/rbt.v57i3.5476

Johow, F. (1911). Observaciones sobre los Onicóforos Chilenos. Boletín Museo Nacional de Historia Natural, 3, 79-98.

Kirk, T.W. (1883). Habitat of Peripatus novo-zealandie. New Zealand Journal of Science, 1, 573.

Krishnan, G. (1970). Chemical nature of the cuticle and its mode of hardening in Eoperipatus weldoni. Acta Histochemica, 37, 1-17.

Laat, D. (2006). Variabilidade genética e estrutura populacional de Peripatus acacioi na estação ecológica do Tripuí, MG. Retrieved from http://www.sidalc.net/cgi-bin/wxis.exe/?IsisScript=AGB.xis&method=post&formato=2&cantidad=1&expresion=mfn=240896

Lacorte, G. A., De Sena Oliveira, I., & Da Fonseca, C. G. (2011). Phylogenetic relationships among the Epiperipatus lineages (Onychophora: Peripatidae) from the Minas Gerais State, Brazil. Zootaxa, 2755, 57-65. https://doi.org/10.11646/zootaxa.2755.1.3

Lankester, E. R. (1904). On the movements of the parapodia of Peripatus, millipedes and centipedes. Quarterly Journal of Microscopical Science, 47, 577-582.

Lavallard, R. (1965). Étude au microscope électronique de l'épithélium tégumentaire chez Peripatus acacioi Marcus et Marcus. Comptes Rendus Hebdomadaires des Seances de l'Academie des Sciences, 260, 965-968.

Lavallard, R., Campiglia, S., Parisi Alvares, E., & Valle, C. M. C. (1975). Contribution a la biologie de Peripatus acacioi Marcus et Marcus (Onychophore). III. Etude descriptive de l'habitat. Vie et Milieu, 25, 87-118.

Lawrence, R. F. (1950). Peripatus - A living museum of antiquities. African Wildlife, 4, 112-120.

Lawrence, R. F. (1977). Insects, arachnids and Peripatus. In A. C. Brown (Ed.), A History of Scientific Endeavour in South Africa (pp. 109-131). Cape Town, South Africa: Royal Society of South Africa.

Locke, M. & Huie, P. (1977). Bismuth staining of Golgi complex is a characteristic arthropod feature lacking in Peripatus. Nature, 270, 341-343. https://doi.org/10.1038/270341a0

Manton, S. M. (1937). Studies on the Onychophora. IV. Proceedings of the Royal Society of London, Part B, 124, S41.

Manton, S. M. (1972). The evolution of arthropodan locomotory mechanisms. Part 10. Locomotory habits, morphology and evolution of the hexapod classes. Zoological Journal of the Linnean Society, 51, 203-400. https://doi.org/10.1111/j.1096-3642.1972.tb02550.x

Marcus, E. (1937). Sobre os Onychophoros. Arquivos do Instituto Biologico, 8, 255-266.

Mayer, G., & Whitington, P. M. (2009). Velvet worm development links myriapods with chelicerates. Proceedings of the Royal Society B: Biological Sciences, 276(1673), 3571-3579. DOI: 10.1098/rspb.2009.0950

Mendes, E. G., & Sawaya, P. (1958). The oxygen consumption of "Onychophora" and its relation to size, temperature and oxygen tension. Revista Brasileira de Biologia, 18, 129-142.

Mesibov, B. (1998). Curious, yes, but not all that rare. Invertebrata, 11, 6.

Milne, L. J., & Milne, M. J. (1954). The worm that didn't turn into anything (Peripatus). Natural History [New York, USA], 63, 182-187.

Monge-Nájera, J. (1994a). Ecological biogeography in the phylum Onychophora. Biogeographica, 70, 111-123.

Monge-Nájera, J. (1994b). Reproductive trends, habitat type and body characteristics in velvet worms (Onychophora). Revista de Biología Tropical, 42, 611-622.

Monge-Nájera, J. (1995). Phylogeny, biogeography and reproductive trends in the Onychophora. Zoological Journal of the Linnean Society, 114, 21-60. https://doi.org/10.1111/j.1096-3642.1995.tb00111.x

Monge-Nájera, J. (1996). Jurassic-Pliocene biogeography: Testing a model with velvet worm (Onychophora) vicariance. Revista de Biología Tropical, 44, 159-175.

Monge-Nájera, J. (2018). City Worms (Onychophora): why do fragile invertebrates from an ancient lineage live in heavily urbanized areas? UNED Research Journal, 10(1), 91-94. DOI: 10.22458/urj.v10i1.2045

Monge-Nájera, J. (2019). "I, astonished, discovered by chance the only specimen”: the first velvet worm (Onychophora). Columna Darwin In Memoriam, Revista de Biología Tropical, Universidad de Costa Rica. Recuperado de https://revistas.ucr.ac.cr/index.php/rbt/article/view/39056

Monge-Nájera, J., & Alfaro, J. P. (1995). Geographic variation of habitats in Costa Rican velvet worms (Onychophora: Peripatidae). Biogeographica, 71, 97-108.

Monge-Nájera, J., & Hou, X. (2000). Disparity, decimation and the Cambrian “explosion”: Comparison of early Cambrian and Present faunal communities with emphasis on velvet worms (Onychophora). Revista de Biologia Tropical, 48(2-3), 333-351.

Monge-Nájera, J., & Lourenco, W. R. (1995). Biogeographic implications of evolutionary trends in onychophorans and scorpions. Biogeographica, 71, 179-185.

Monge-Nájera, J., & Morera-Brenes, B. (2015). Velvet Worms (Onychophora) in Folklore and Art: Geographic Pattern, Types of Cultural Reference and Public Perception. British Journal of Education, Society and Behavioural Science, 10(3), 1-9. https://doi.org/10.9734/BJESBS/2015/18945

Monge-Nájera, J., & Xianguang, H. (2002). Experimental taphonomy of velvet worms (Onychophora) and implications for the Cambrian “explosion, disparity and decimation” model. Revista de Biologia Tropical, 50(3-4), 1133-1138.

Monge-Nájera, J., Barquero-González, P., & Morera-Brenes, B. (2019a). Why do velvet worm spermatozoa swim for years? Columna Darwin In Memoriam, Revista de Biología Tropical, Universidad de Costa Rica. Recuperado de https://revistas.ucr.ac.cr/index.php/rbt/article/view/36240

Monge-Nájera, J., Barquero-González, P., & Morera-Brenes, B. (2019b). Two ways to be a velvet worm. Columna Darwin In Memoriam, Revista de Biología Tropical, Universidad de Costa Rica. Recuperado de https://revistas.ucr.ac.cr/index.php/rbt/article/view/36107

Monge-Nájera, J., Barquero-González, P., & Morera-Brenes, B. (2019c). Why are there no onychophorans in Cuba? Columna Darwin In Memoriam, Revista de Biología Tropical, Universidad de Costa Rica. Recuperado de https://revistas.ucr.ac.cr/index.php/rbt/article/view/36418

Monge-Nájera, J., Barquero-González, P., & Morera-Brenes, B. (2019d). Why do onychophorans and camels need the same gestation time? Columna Darwin In Memoriam, Revista de Biología Tropical, Universidad de Costa Rica. Recuperado de https://revistas.ucr.ac.cr/index.php/rbt/article/view/36118

Monge-Nájera, J., Barquero-González, P., & Morera-Brenes, B. (2019e). Are the eggs - kept and fed by the mother - the ancestral form of reproduction in onychophorans? Columna Darwin In Memoriam, Revista de Biología Tropical, Universidad de Costa Rica. Recuperado de https://revistas.ucr.ac.cr/index.php/rbt/article/view/36398

Monge-Nájera, J., Barquero-González, P., & Morera-Brenes, B. (2019f). What do we really know about how velvet worms mate? Columna Darwin In Memoriam, Revista de Biología Tropical, Universidad de Costa Rica. Recuperado de https://revistas.ucr.ac.cr/index.php/rbt/article/view/36337

Monge-Nájera, J., Barquero-González, P., & Morera-Brenes, B. (2019g). Why do some Australian onychophorans have fantastic heads? Columna Darwin In Memoriam, Revista de Biología Tropical, Universidad de Costa Rica. Recuperado de https://revistas.ucr.ac.cr/index.php/rbt/article/view/36337

Monge-Nájera, J., Barrientos, Z., & Aguilar, F. (1993). Behavior of Epiperipatus biolleyi (Onychophora: Peripatidae) under laboratory conditions. Revista de Biología Tropical, 41, 689-696.

Montgomery, T. H. (1900). The spermatogenesis of Peripatus (Peripatopsis) balfouri up to the formation of the spermatid. Zoologische Jahrbücher, Abteilung für Anatomie und Ontogenie der Tiere, 14, 277-368.

Montgomery, T. H. (1912). Complete discharge of mitochondria from the spermatozoon of Peripatus. Biological Bulletin, 22, 309319. https://doi.org/10.2307/1535903

Mora, M., Herrera, A. & León, P. (1996). Analisis electroforético de las secreciones adhesivas de onicoforos del género Epiperipatus (Onychophora: Peripatidae). Revista de Biología Tropical, 44, 147-152.

Morera, B., Monge-Nájera, J., & Mora, P. C. (2018). The Conservation Status of Costa Rican Velvet Worms (Onychophora). DOI: 10.20944/PREPRINTS201812.0151.V1

Morera, B., Monge-Nájera, J., & Saenz, R. (1988). Parturition in onychophorans: New record and a review. Brenesia, 29, 15-20.

Morera-Brenes, B., & Monge-Nájera, J. (2010). A new giant species of placented worm and the mechanism by which onychophorans weave their nets (Onychophora: Peripatidae). Revista de Biologia Tropical, 58(4), 1127-1142. https://doi.org/10.15517/rbt.v58i4.5398

Morera-Brenes, B., Herrera, A., Mora, M. & León, P. (1992). Estudios genómicos de Epiperipatus biolleyi (Peripatidae, Onychophora). Revista Brasileira de Genética (Brazilian Journal of Genetics), 15: 91.

Morris, S. C. (1977). A new metazoan from the Cambrian Burgess Shale of British Columbia. Palaeontology, 20(3), 623-40.

Nicolas, A. (1889). Sur les rapports des muscles et des éléments épithéliaux dans le pharynx du Péripate (Peripatus capensis). Revue Biologique du Nord de la France, 2, 81-98

Oliveira, I. de S.; Read, V. M. S. J., & Mayer, G. (2012). A world checklist of onychophora (velvet worms), with notes on nomenclature and status of names. ZooKeys, 211, 1-70. https://doi.org/10.3897/zookeys.211.3463

Packard, A. S. (1898). Relations of Peripatus to insects. In A. S. Packard (Ed.), A text-book of Entomology (pp. 9-11). New York: The MacMillan Company.

Pass, G. (1991). Antennal circulatory organs in Onychophora, Myriapoda and Hexapoda: Functional morphology and evolutionary implications. Zoomorphology, 110, 145-164. https://doi.org/10.1007/BF01632871

Peck, S. B. (1975). A review of the New World Onychophora with the description of a new cavernicolous genus and species from Jamaica. Psyche, 82, 341-358. https://doi.org/10.1155/1975/98614

Periera, J., Bicudo, W., & Campiglia, S. (1985). A morphometric study of the tracheal system of Peripatus acacioi Marcus and Marcus (Onychophora). Respiration Physiology, 60(1), 75-82. https://doi.org/10.1016/0034-5687(85)90040-4

Peters, W. (1880). Die Variation der Fusszahl bei Peripatus capensis Grube. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin [1880], 165-166.

Picado, C. (1911). Sur un habitat nouveau de Peripatus. Bulletin du Museum National d'Historie Naturelle, 17, 415-416.

Poinar, G. (1996). Fossil velvet worms in Baltic and Dominican amber: Onychophoran evolution and biogeography. Science, 273, 1370-1371.

Poinar, Jr. G. (2000). Fossil onychophorans from Dominican and Baltic amber: Tertiapatus dominicanus n.g., n.sp. (Tertiapatidae n.fam.) and Succinipatopsis balticus n.g., n.sp. (Succinipatopsidae n.fam.) with a proposed classification of the subphylum Onychophora. Invertebrate Biology, 119(1), 104-109. DOI: 10.1111/j.1744-7410.2000.tb00178.x

Porter, C. E. (1905). Lecciones de historia natural dictadas a los alumnos del 4. año de la Escuela Naval (curso de 1904): Los Onicóforos. Revista Chilena de Historia Natural ,9, 124-128.

Porter, C. E. (1917). Bibliografía chilena razonada de Miriápodos y Onicóforos. Revista Chilena de Historia Natural, 21, 52-62.

Prenant, A. (1890). Note sur les éléments séminaux d'un Peripatus. Revue Biologique du Nord de la France, 2, 169-174.

Read, J. (1988). The application of scanning electron microscopy to the systematics of the neotropical Peripatidae (Onychophora). Zoological Journal of the Linnean Society, 93, 187-223. https://doi.org/10.1111/j.1096-3642.1988.tb01361.x

Reinhard, J., & Rowell, D. M. (2005). Social behaviour in an Australian velvet worm, Euperipatoides rowelli (Onychophora: Peripatopsidae). Journal of Zoology, 267(1), 1-7. https://doi.org/10.1017/S0952836905007090

Rhebergen, F., & Donovan, S. K. (1994). A lower Palaeozoic "onychophoran" reinterpreted as a pelmatozoan (stalked echnioderm) column. Atlantic Geology, 30, 19-23. https://doi.org/10.4138/2116

Rolfe, W. D. I.; Schram, F. R.; Pacaud, G.; Sotty, D., & Secretan, S. (1982). A remarkable Stephanian biota from Montceau-les-Mines, France. Journal of Paleontology, 56, 426-428.

Ruhberg, H. (1979). Light and electron microscopical investigations on the salivary glands of Opisthopatus cinctipes and Peripatopsis moseleyi (Onychophora: Peripatopsidae). Zoologischer Anzeiger, 203, 35-47.

Ruhberg, H. (1985). Die Peripatopsidae (Onychophora). Systematik, Ökologie, Chorologie und phylogenetische Aspekte. In F. Schaller (Ed.), Zoologica. Stuttgart: Schweizerbart'sche Verlagsbuchhandlung.

Ruhberg, H.; Tait, N. N.; Briscoe, D. A. & Storch, V. (1988). Cephalofovea tomahmontis n. gen., n. sp., an Australian peripatopsid (Onychophora) with a unique cephalic pit. Zoologischer Anzeiger, 221, 117-133.

Ryu, S. H.; Kang, C. W.; Choi, J.; Myung, Y.; Ko, Y.J.; Lee, S. M.; Son, S. U. (2018). Microporous Porphyrin Networks Mimicking a Velvet Worm Surface and Their Enhanced Sensitivities toward Hydrogen Chloride and Ammonia. ACS Applied Materials & Interfaces, 10(8), 6815-6819. DOI: 10.1021/acsami.7b19119

Saint-Remy, G. (1889). Sur la structure du cerveau chez les Myriopodes et les Arachnides. Revue Biologique du Nord de la France, 2, 41-55.

Sanchez, S. (1958). Cellules neurosécrétrices et organes infracérébraux de Peripatopsis moseleyi Wood (Onychophores) et neurosécrétion chez Nymphon gracile Leach (Pycnogonides). Archives de Zoologie Experimentale et Generale [Notes et Revue], 96, 57-62.

Santana, G. G., Almeida, W. D. O., Alves, R. R. da N., & Vasconcellos, A. (2008). Extension of the northern distribution of Onychophora in the Brazilian Atlantic Forest. Biotemas, 21(2), 161-163. DOI: 10.5007/2175-7925.2008v21n2p161

Schürmann, F. W., & Sandeman, D. C. (1976). Giant fibres in the ventral nerve cord of Peripatoides leuckarti (Onychophora). Naturwissenschaften, 63, 580-581. https://doi.org/10.1007/BF00622801

Sclater, W. L. (1888). On the early stages of the development of South American species of Peripatus. Quarterly Journal of Microscopical Science, 28, 343-363.

Sedgwick, A. (1884). On the origin of metameric segmentation and some other morphological questions. Quarterly Journal of Microscopical Science, 24, 43-82.

Sedgwick, A. (1908a). The distribution and classification of the Onychophora. Quarterly Journal of Microscopical Science, 52, 379-406.

Sedgwick, A. (1908b). Relation between the geographical distribution and the classification of the Onychophora. Proceedings of the Cambridge Philosophical Society, 14, 546.

Sheldon, L. (1887). On the development of Peripatus novae-zealandiae. Quarterly Journal of Microscopical Science, 28, 205-237.

Sheldon, L. (1890). The maturation of the ovum in the Cape and New Zealand species of Peripatus. Quarterly Journal of Microscopical Science, 30, 1-29

Silva, J. R. M. C. da, Coelho, M. P. D., & Nogueira, M. I. (2000). Processo inflamatório induzido em Peripatus acacioi (onychophora) um fóssil vivo. Journal Invertebrate Pathology, 75, 41-46. https://doi.org/10.1006/jipa.1999.4898

Simoes, L. C. G., Marques da Silva, I., & Schreiber, G. (1964). DNA e volume nuclear em tecidos de Peripatus acacioi (Marcus & Marcus). Ciencia e Cultura [Sao Paulo], 16, 291-295.

Snodgrass, R. E. (1938). Evolution of the Annelida, Onychophora and Arthropoda. Smithsonian Miscellaneous Collections, 97, 1-159.

Sosa-Bartuano, Á., Monge-Nájera, J., & Morera-Brenes, B. (2018). A proposed solution to the species problem in velvet worm conservation (Onychophora). UNED Research Journal, 10(1), 204-208. https://doi.org/10.22458/urj.v10i1.2027

Storch, V., & Ruhberg, H. (1977). Fine structure of the sensilla of Peripatopsis moseleyi (Onychophora). Cell & Tissue Research, 177, 539-553. https://doi.org/10.1007/BF00220613

Storch, V.; Alberti, G.; Lavallard, R., & Campiglia, S. S. (1979). Données ultrastructurales sur des zones synaptiques dans les nephridies de Peripatus acacioi Marcus et Marcus (Onychophore). Boletim de Fisiologia Animal, 3, 145-156.

Tait, N. N., & Norman, J. M. (2001). Novel mating behaviour in Florelliceps stutchburyae gen. nov., sp. nov. (Onychophora: Peripatopsidae) from Australia. Journal of Zoology, 253(3), 301-30. https://doi.org/10.1017/S0952836901000280

Tarlo, L. B. (1967). Xenusion - Onychophoran or Coelenterate? Mercian Geologist, 2, 97-99.

Thompson, I., & Jones, D. S. (1980). A possible onychophoran from the Middle Pennsylvanian Mazon Creek Beds of northern Illinois. Journal of Paleontology, 54, 588-596.

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(5), 403-416. DOI: 10.1002/dvdy.24492

Trindade, G. (1958). Peripatus e sua possivel utilizacao em laboratorio. Anais do III Congresso de Farmacia e Bioquimica Pan-americano e V Congresso Brasileiro de Farmacia 519-520.

Tuzet, O., & Manier, J. F. (1958). Recherches sur Peripatopsis moseleyi Wood-Mason péripate du Natal. I. Étude sur le sang. II. La spermatogenèse. Bulletin Biologique de la France et de la Belgique, 92, 7-23.

Vachon, M. (1954). Répartition actuelle et ancienne des Onychophores ou Peripates. Revue Générale des Sciences Pures et Appliquées et Bulletin de l'Association Francaise pour l'Avancement des Sciences, 61, 300-308.

Van der Lande, V.M. (1991). Native and introduced Onychophora in Singapore. Zoological Journal of the Linnean Society, 102, 101-114. https://doi.org/10.1111/j.1096-3642.1991.tb00284.x

Vincent, M. (1936). Un nouveau type de Coccidie des Péripates. Comptes Rendus des Seances de la Société de Biologie et de ses Filiales, 122, 260-262.

Wägele, J. W. (1993). Rejection of the "Uniramia" hypothesis and implications of the Mandibulata concept. Zoologische Jahrbücher, Abteilung für Systematik, Ökologie und Geographie der Tiere, 120, 253-288.

Walcott, C. D. (1931). Addenda to descriptions of Burgess shale fossils. Smithsonian Miscellaneous Collections, 85, 1-46.

Wells, S. M.; Pyle, R. M., & Collins, N. M. (1983). Peripatus. Phylum Onychophora. In The IUCN Invertebrate Red Data Book IUCN (pp. 515-520). Gland, Switzerland.

Wenzel, R. L. (1950). Peripatus - "living fossil" and "missing link". Tuatara, 3, 98-99.

Whitman, C.O. (1891). Spermatophores as a means of hypodermic impregnation. Journal of Morphology, 4, 361-406. https://doi.org/10.1002/jmor.1050040307

Wood-Mason, J. (1879). Morphological notes bearing on the origin of insects. Transactions of the Entomological Society of London [1879], 2, 145-167.

Zilch, A. (1955). Begegnung mit Peripatus. Natur und Volk, 85, 146-154.



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Onychophorology, the study of velvet worms, historical trends, landmarks, and researchers from 1826 to 2020 (a literature review). (2021). Uniciencia, 35(1), 210-230. https://doi.org/10.15359/ru.35-1.13



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How to Cite

Onychophorology, the study of velvet worms, historical trends, landmarks, and researchers from 1826 to 2020 (a literature review). (2021). Uniciencia, 35(1), 210-230. https://doi.org/10.15359/ru.35-1.13

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