A new giant velvet worm from Costa Rica suggests absence of the genus Peripatus (Onychophora: Peripatidae) in Central America

Authors

  • José Pablo Barquero González Universidad Nacional de Costa Rica Author http://orcid.org/0000-0001-8343-6050
  • Steven Sánchez-Vargas Universidad Nacional Author
  • Bernal Morera-Brenes Universidad Nacional Author

DOI:

https://doi.org/10.15517/rbt.v68i1.37675

Keywords:

onychophoran speciation, Costa Rican biodiversity, endemism, saproxilic taxa, phylogenetics

Abstract

Introduction: Neotropical onychophoran taxonomy and diversity has been poorly investigated. Recent studies have discovered problems in species classification: they have questioned the accepted genera and the actual number of species. This is true in Costa Rica, where several unidentified species have been reported. Objective: The objective of this investigation was to evaluate the occurrence of the accepted genera in this country, and to describe a new genus and species from Central America. Methods: In 2017, we collected one onychophoran in the Keköldi Indigenous Reserve in Talamanca, Limón, Costa Rica. The specimen gave birth to several offspring. Therefore, seven organisms were analyzed. Light microscopy was used to observe the gross morphology in all samples. The detailed morphology was studied in the biggest specimen with scanning electron microscopy; after that, we performed a phylogenetic analysis with the corresponding sequence of COI. Results: According to our results, a new genus and species of giant onychophoran was found. The genus was identified by its giant size, apical piece of seven scale ranks, large conical primary papillae, dorso-median furrow flanked by two-three accessory papillae, the absence of hyaline organs and a marked sexual dimorphism with respect to the number of legs. The new species presents a particular head pattern, as well as novel structures like cephalic papillae, accessory papillae with rudimentary apical pieces, and a lack of antennal chemoreceptors. Phylogenetic analysis rendered our genus as monophyletic and includes Peripatus solorzanoi, which is grouped within the Central American clade. As our species is clustered inside the Costa Rica-Panamanian group, it is not related to the Caribbean Island nor Guyanan Shield samples, home of Epiperipatus and Peripatus respectively. Therefore, we suggest that those genera do not occur in Central America, and a new genus exists: Mongeperipatus, gen. nov. Conclusion: We concluded that Costa Rica is home to a diversity of undescribed onychophorans that requires specific studies to help clarify the taxonomy and evolutionary relationships of the group to justify their protection.

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Author Biographies

  • José Pablo Barquero González, Universidad Nacional de Costa Rica
    Escuela de Ciencias Biológicas.Laboratorio de Sistemática,Genética y Evolución (LabSGE). Universidad Nacional de Costa Rica.https://sites.google.com/site/labgeweb/home Investigator and student.
  • Steven Sánchez-Vargas, Universidad Nacional

    Escuela de Ciencias Biológicas

  • Bernal Morera-Brenes, Universidad Nacional

    Escuela de Ciencias Biológicas

References

Akaike, H. (1973). Maximum likelihood identification of Gaussian autoregressive moving average models. Biometrika, 60(2), 255-265. DOI: https://doi.org/10.1093/biomet/60.2.255

Allwood, J., Gleeson, D., Mayer, G., Daniels, S., Beggs, J.R., & Buckley, T.R. (2010). Support for vicariant origins of the New Zealand Onychophora. Journal of Biogeography, 37(4), 669-681. DOI: https://doi.org/10.1111/j.1365-2699.2009.02233.x

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: https://doi.org/10.1021/acs.biomac.8b01017

Barquero-González, J.P., Acosta-Chaves, V.J., Sotela, M.L., Brenes, F.V., & Morera-Brenes, B. (2016a). Evidencia fotográfica de especies desconocidas de onicóforos (Onychophora: Peripatidae) de Costa Rica. UNED Research Journal, 8(2), 139-147. DOI: https://doi.org/10.22458/urj.v8i2.1553

Barquero-González, J.P., Alvarado Cabrera, A., Valle-Cubero, S., Monge-Nájera, J., & Morera-Brenes, B. (2016b). The geographic distribution of Costa Rican velvet worms (Onychophora: Peripatidae). Revista de Biología Tropical, 64(4), 1401-1414. DOI: https://doi.org/10.15517/rbt.v64i4.19486

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. DOI: 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: https://doi.org/10.22458/urj.v11i2.2195

Barnes, A., & Daniels, S.R. (2019). On the importance of fine-scale sampling in detecting alpha taxonomic diversity among saproxylic invertebrates: A velvet worm (Onychophora: Opisthopatus amaxhosa) template. Zoologica Scripta, 48(2), 243-262. DOI: https://doi.org/10.1111/zsc.12338

Bouvier, E.L. (1899). Contribution a l’histoire des Péripates Américanes. Annales de la Societé Entomologique de France, 68, 385-450. DOI: https://doi.org/10.1080/21686351.1899.12279742

Bouvier, E.L. (1902a). Un Péripate de Costa-Rica. Bulletin de la Société Entomologique de France, 7(15), 239-240. DOI: https://doi.org/10.3406/bsef.1902.23111

Bouvier, E.L. (1902b). Peripatus biolleyi, Onychophore nouveau de Costa Rica. Bulletin de la Société Entomologique de France, 16, 258-259. DOI: https://doi.org/10.3406/bsef.1902.23123

Bouvier, E.L. (1905). Monographie des Onychophores I. Annales des Sciences Naturelles, 9(2), 1-383.

Braband, A., Podsiadlowski, L., Cameron, S.L., Daniels, S., & Mayer, G. (2010a). Extensive duplication events account for multiple control regions and pseudo-genes in the mitochondrial genome of the velvet worm Metaperipatus inae (Onychophora, Peripatopsidae). Molecular Phylogenetics and Evolution, 57(1), 293-300. DOI: https://doi.org/10.1016/j.ympev.2010.05.012

Braband, A., Cameron, S.L., Podsiadlowski, L., Daniels, S.R., & Mayer, G. (2010b). The mitochondrial genome of the onychophoran Opisthopatus cinctipes (Peripatopsidae) reflects the ancestral mitochondrial gene arrangement of Panarthropoda and Ecdysozoa. Molecular Phylogenetics and Evolution, 57(1), 285-292. DOI: https://doi.org/10.1016/j.ympev.2010.05.011

Brinck, P. (1957). Onychophora, a review of South African species, with a discussion on the significance of the geographical distribution of the group. South African Animal Life, 4, 7-32.

Brito, S.V., Pereira, J.C., Ferreira, F.S., Vasconcellos, A., & Almeida, W.O. (2010). Epiperipatus cratensis sp. nov. (Onychophora: Peripatidae) from northeastern Brazil. Neotropical Biology and Conservation, 5(1), 47-52. DOI: https://doi.org/10.4013/nbc.2010.51.07

Briscoe, D.A., & Tait, N.N. (1995). Allozyme evidence for extensive and ancient radiations in Australian Onychophora. Zoological Journal of the Linnean Society, 114(1), 91-102. DOI: https://doi.org/10.1111/j.1096-3642.1995.tb00114.x

Castresana, J. (2000). Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Molecular Biology and Evolution, 17(4), 540-552. DOI: https://doi.org/10.1093/oxfordjournals.molbev.a026334

Carvalho, A. (1941). Nota prévia sôbre uma nova espécie de “Peripatus” do Brasil Central. Brazilian Journal of Biology, 1(4), 447-448.

Chagas-Júnior, A., & Sampaio Costa, C. (2014). Macroperipatus ohausi: redescription and taxonomic notes on its status (Onychophora: Peripatidae). International Journal of Tropical Biology and Conservation, 62(3), 977-985. DOI: https://doi.org/10.15517/rbt.v62i3.11643

Clark, A.H. (1913). A revision of the American species of Peripatus. Proceedings of the Biological Society of Washington, 26, 15-19.

Cockerell, T.D.A. (1908). Monographie des Onychophores by Bouvier. Science, 27, 619-621. DOI: https://doi.org/10.1126/science.27.694.619

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

Corrales-Ureña, 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), 125013. DOI: https://doi.org/10.1088/2053-1591/aa9940

Cunha, W.T., Santos, R.C., 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: https://doi.org/10.1590/1678-4685-gmb-2016-0037

Daniels, S.R. (2011). Genetic variation in the Critically Endangered velvet worm Opisthopatus roseus (Onychophora: Peripatopsidae). African Zoology, 46(2), 419-424. DOI: https://doi.org/10.3377/004.046.0222

Daniels, S.R., Picker, M.D., Cowlin, R.M., & Hamer, M.L. (2009). Unravelling evolutionary lineages among South African velvet worms (Onychophora: Peripatopsis) provides evidence for widespread cryptic speciation. Biological Journal of the Linnean Society, 97(1), 200-216. DOI: https://doi.org/10.1111/j.1095-8312.2009.01205.x

Daniels, S.R., & Ruhberg, H. (2010). Molecular and morphological variation in a South African velvet worm Peripatopsis moseleyi (Onychophora, Peripatopsidae): evidence for cryptic speciation. Journal of Zoology, 282(3), 171-179. DOI: https://doi.org/10.1111/j.1469-7998.2010.00722.x

Daniels, S.R., McDonald, D.E., & Picker, M.D. (2013). Evolutionary insight into the Peripatopsis balfouri sensu lato species complex (Onychophora: Peripatopsidae) reveals novel lineages and zoogeographic patterning. Zoologica Scripta, 42(6), 656-674. DOI: https://doi.org/10.1111/zsc.12025

Daniels, S.R., Dambire, C., Klaus, S., & Sharma, P.P. (2016). Unmasking alpha diversity, cladogenesis and biogeographical patterning in an ancient panarthropod lineage (Onychophora: Peripatopsidae: Opisthopatus cinctipes) with the description of five novel species. Cladistics, 32(5), 506-537. DOI: https://doi.org/10.1111/cla.12154

Dzik, J., & Krumbiegel, G. (1989). The oldest onychophoran, Xenusion - a link connecting phyla. Lethaia, 22, 169-181. DOI: https://doi.org/10.1111/j.1502-3931.1989.tb01679.x

Evans, R. (1901). On two new species of Onychophora from the Siamese Malay States. Quarterly Journal of Microscopical Science, 44, 473-538. DOI: https://doi.org/10.1242/jcs.s2-44.176.473

Fuhrmann, O. (1913). Quelques nouveaux Péripates américains. Mémoires de la Société Neuchâteloise des Sciences Naturelles, 5, 176-192.

Garwood, R.J., Edgecombe, G.D., Charbonnier, S., Chabard, D., Sotty, D., & Giribet, G. (2016). Carboniferous Onychophora from Montceau les Mines, France, and onychophoran terrestrialization. Invertebrate Biology, 135(3), 179-190. DOI: https://doi.org/10.1111/ivb.12130

Giribet, G., Buckman-Young, R.S., Costa, C.S., Baker, C.M., Benavides, L.R., Branstetter, M.G., & Pinto-da-Rocha, R. (2018). The ‘Peripatos’ in Eurogondwana?–Lack of evidence that south-east Asian onychophorans walked through Europe. Invertebrate Systematics, 32(4), 840-863. DOI: https://doi.org/10.1071/IS18007

Gleeson, D.M., & Ruhberg, H. (2010). Phylum Onychophora–velvet worms, peripatus. New Zealand Inventory of Biodiversity: Kingdom Animalia, 2, 36-39.

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

Haritos, V.S., Niranjane, A., Weisman, S., Trueman, H.E., Sriskantha, A., & Sutherland, T.D. (2010). Harnessing disorder: onychophorans use highly unstructured proteins, not silks, for prey capture. Proceedings of the Royal Society of London B: Biological Sciences, 277(1698), 3255-3263. DOI: https://doi.org/10.1098/rspb.2010.0604

Holdridge, L.R. (1967). Life zone ecology. San José, Costa Rica: Centro Científico Tropical.

Katoh, K., Kuma, K., Toh, H., & Miyata, T. (2005). MAFFT version 5: improvement in accuracy of multiple sequence alignment. Nucleic Acids Research, 33, 511-518. DOI: https://doi.org/10.1093/nar/gki198

Kemp, S. (1913). Preliminary note on a new genus of Onychophora from the N. E. frontier of India. Records of the Indian Museum, 9, 241-242.

von Kennel, J. (1883). Entwicklungsgeschichte von Peripatus. Zoologischer. Anzeiger, 6(150), 531-537.

Kuraku, S., Zmasek, C.M., Nishimura, O., & Katoh, K. (2013). aLeaves facilitates on-demand exploration of metazoan gene family trees on MAFFT sequence alignment server with enhanced interactivity. Nucleic Acids Research, 41, W22-W28. DOI: https://doi.org/10.1093/nar/gkt389

Lacorte, G.A., Oliveira, I.S., & Fonseca, C.G. (2011). Population structure and demographic inferences concerning the endangered onychophoran species Epiperipatus acacioi (Onychophora: Peripatidae). Genetics and Molecular Research, 10(4), 2775-2785. DOI: https://doi.org/10.4238/2011.November.9.1

Lavallard, R., & Campiglia, S. (1975). Contribution a la biologie de Peripatus acacioi Marcus et Marcus (Onychophore). V. Etude des Naissances dans un elevage de laboratiore. Zoologischer Anzeiger, 195, 338-350.

Marcus, E., & Marcus, E. (1955). A new Peripatus from Minas Gerais, Brazil. Anais da Academia Brasileira de Ciencias, 27, 189-193.

Mayer, G. (2007). Metaperipatus inae sp. nov. (Onychophora: Peripatopsidae) from Chile with a novel ovarian type and dermal insemination. Zootaxa, 1440(1), 21-37. DOI: https://doi.org/10.11646/zootaxa.1440.1.2

Miller, M.A., Pfeiffer, W., & Schwartz, T. (2010). Creating the CIPRES Science Gateway for inference of large phylogenetic trees. Gateway Computing Environments Workshop, 1-8. DOI: https://doi.org/10.1109/GCE.2010.5676129

Monge-Nájera, J. (1994). Reproductive trends, habitat type and body characteristics in velvet worms (Onychophora). Revista de Biología Tropical, 42(3), 611-622.

Monge-Nájera, J. (1995). Phylogeny, biogeography and reproductive trends in the Onychophora. Zoological Journal of the Linnean Society, 114(1), 21-60. DOI: https://doi.org/10.1006/zjls.1995.0015

Monge-Nájera, J. (1996). Jurassic–Pliocene biogeography: testing a model with velvet worm (Onychophora) vicariance. International Journal of Tropical Biology and Conservation, 44(1), 147-152.

Monge-Nájera, J. (2017). The power of short lectures to improve support for biodiversity conservation of unpopular organisms: an experiment with worms. UNED Research Journal, 9(1), 145-150. DOI: https://doi.org/10.22458/urj.v9i1.1690

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: https://doi.org/10.22458/urj.v10i1.2045

Monge-Nájera, J., Barrientos, Z., & Aguilar, F. (1993). Behavior of Epiperipatus biolleyi (Onychophora: Peripatidae) under laboratory conditions. Revista de Biología Tropical, 41(3A), 689-696.

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 & Behavioural Science, 10(3), 1-9. DOI: https://doi.org/10.9734/BJESBS/2015/18945

Mora, M., Herrera, A., & León, P. (1996). The genome of Epiperipatus biolleyi (Peripatidae), a Costa Rican onychophoran. Revista de Biología Tropical, 44(1), 153-157.

Morera-Brenes, B., & León, M.D. (1986). Description of Macroperipatus valerioi n. sp. from Costa Rica, and comments on the genus Macroperipatus (Onychophora: Peripatidae). Revista de Biología Tropical, 34(2), 277-282.

Morera Brenes, B., Monge-Nájera, J., & Sáenz, R. (1988). Parturition in Onychophorans: New Record and a Review. Brenesia, 29, 15-20.

Morera-Brenes, B., & Monge-Nájera, J. (1990). Epiperipatus hilkae, n. sp. from Costa Rica (Onychophora: Peripatidae). Revista de Biología Tropical, 38(2B), 449-456.

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 Biología Tropical, 58(4), 1127-1142. DOI: https://doi.org/10.15517/rbt.v58i4.5398

Murienne, J., Daniels, S.R., Buckley, T.R., Mayer, G., & Giribet, G. (2014). A living fossil tale of Pangaean biogeography. Proceedings of the Royal Society of London B: Biological Sciences, 281(1775), 20132648. DOI: https://doi.org/10.1098/rspb.2013.2648

Myburgh, A.M., & Daniels, S.R. (2015). Exploring the impact of habitat size on phylogeographic patterning in the Overberg velvet worm Peripatopsis overbergiensis (Onychophora: Peripatopsidae). Journal of Heredity, 106(3), 296-305. DOI: https://doi.org/10.1093/jhered/esv014

Oliveira, I.D.S., & Wieloch, A.H. (2005). Macroperipatus machadoi sp. n. (Onychophora: Peripatidae) da floresta atlântica de Minas Gerais, Brasil. Lundiana, 6(supplement), 61-66. DOI: https://doi.org/10.35699/2675-5327.2005.22120

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. DOI: https://doi.org/10.11646/zootaxa.2493.1.2

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. DOI: https://doi.org/10.1371/journal.pone.0019973

Oliveira, I.S., Franke, F.A., Hering, L., Schaffer, S., Rowell, D.M., Weck-Heimann, A., … Mayer, G. (2012a). Unexplored character diversity in Onychophora (velvet worms): a comparative study of three peripatid species. PloS One, 7(12), e51220. DOI: https://doi.org/10.1371/journal.pone.0051220

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

Oliveira, I.S., Schaffer, S., Kvartalnov, P.V., Galoyan, E.A., Plako, 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-A Journal of Comparative Zoology, 252(4), 495-510. DOI: https://doi.org/10.1016/j.jcz.2013.01.001

Oliveira, I.S., Lacorte, G.A., Weck-Heimann, A., Cordeiro, L.M., Wieloch, A.H., & Mayer, G. (2014). A new and critically endangered species and genus of Onychophora (Peripatidae) from the Brazilian savannah–a vulnerable biodiversity hotspot. Systematics and Biodiversity, 13(3), 211-233. DOI: https://doi.org/10.1080/14772000.2014.985621

Oliveira, I.S., Bai, M., Jahn, H., Gross, V., Martin, C., Hammel, J.U., … Mayer, G. (2016). Earliest onychophoran in amber reveals Gondwanan migration patterns. Current Biology, 26(19), 2594-2601. DOI: https://doi.org/10.1016/j.cub.2016.07.023

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(4), 909-932. DOI: https://doi.org/10.1071/IS17096

Peck, S.B. (1975). A Review of the New World Onychophora with the Description of a New Cavernicolous Genus and Species from Jamaica. Psyche: A Journal of Entomology, 82(3-4), 341-358. DOI: https://doi.org/10.1155/1975/98614

Picado, C. (1911). Sur un habitat nouveau des Peripatus. Bulletin du Musée National d’Histoire Naturelle, Paris, 17, 415-416.

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(1), 42-51. DOI: https://doi.org/10.1093/molbev/msm223

Posada, D. (2008). jModelTest: phylogenetic model averaging. Molecular Biology and Evolution, 25(7), 1253-1256. DOI: https://doi.org/10.1093/molbev/msn083

Rambaut, A., Drummond, A.J., Xie, D., Baele, G., & Suchard, M.A. (2018). Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Systematic Biology, 67(5), 901-904. DOI: https://doi.org/10.1093/sysbio/syy032

Read, V.S.J. (1985). The ecology of Macroperipatus torquatus (Kennel) with special reference to feeding and a taxonomic review (Doctoral dissertation). University of Wales, United Kingdom.

Read, V.S.J. (1988a). The application of scanning electron microscopy to the systematics of the neotropical Peripatidae (Onychophora). Zoological Journal of the Linnean Society, 93(3), 187-223. DOI: https://doi.org/10.1111/j.1096-3642.1988.tb01361.x

Read, V.S.J. (1988b). The Onychophora of Trinidad, Tobago and the Lesser Antilles. Zoological Journal of the Linnean Society, 93(3), 225-257. DOI: https://doi.org/10.1111/j.1096-3642.1988.tb01362.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. DOI: https://doi.org/10.1017/S0952836905007090

Reid, A.L. (1996). Review of the Peripatopsidae (Onychophora) in Australia, with comments on peripatopsid relationships. Invertebrate Systematics, 10(4), 663-936. DOI: https://doi.org/10.1071/IT9960663

Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., … Huelsenbeck, J.P. (2012). MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61(3), 539-542. DOI: https://doi.org/10.1093/sysbio/sys029

Rota-Stabelli, O., Kayal, E., Gleeson, D., Daub, J., Boore, J.L., Telford, M.J., Pisani, D., Blaxter, M., & Lavrov, D.V. (2010). Ecdysozoan mitogenomics: evidence for a common origin of the legged invertebrates, the Panarthropoda. Genome Biology and Evolution, 2, 425-440. DOI: https://doi.org/10.1093/gbe/evq030

Ruhberg, H. (1985). Die Peripatopsidae (Onychophora), Systematik, Ökologie, Chorologie und phylogenetische Aspekte. Zoologica, 137, 1-184.

Ruhberg, H., & Hamer, M.L. (2005). A new species of Opisthopatus Purcell, 1899 (Onychophora: Peripatopsidae) from KwaZulu-Natal, South Africa. Zootaxa, 1039(1), 27-38. DOI: https://doi.org/10.11646/zootaxa.1039.1.3

Ruhberg, H., & Daniels, S.R. (2013). Morphological assessment supports the recognition of four novel species in the widely distributed velvet worm Peripatopsis moseleyi sensu lato (Onychophora: Peripatopsidae). Invertebrate Systematics, 27(2), 131-145. DOI: https://doi.org/10.1071/IS12069

Sampaio-Costa, C., Chagas-Junior, A., & Baptista, R.L. (2009). Brazilian species of Onychophora with notes on their taxonomy and distribution. Zoologia, 26(3), 553-561. DOI: https://doi.org/10.1590/S1984-46702009005000004

Sampaio-Costa, C. (2016). Sistemática e análise filogenética de Epiperipatus Clark, 1913 baseada em dados moleculares e morfológicos (Onychophora: Peripatidae) (Doctoral dissertation). Universidade de São Paulo, Brasil.

Sampaio-Costa, C., 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: https://doi.org/10.3897/zoologia.35.e23366

Sato, S., Buckman-Young, R.S., Harvey, M.S., & Giribet, G. (2018). Cryptic speciation in a biodiversity hotspot: multilocus molecular data reveal new velvet worm species from Western Australia (Onychophora: Peripatopsidae: Kumbadjena). Invertebrate Systematics, 32(6), 1249-1264. DOI: https://doi.org/10.1071/IS18024

Schmarda, L.K. (1871). I. Ordnung. Malacopoda Blainville. Onychophora. Grube. Zoologie, 1(2), 74-76.

Sedgwick, A. (1888). A monograph on the species and distribution of the genus Peripatus (Guilding). Quarterly Journal of Microscopical Science, 28, 431-493. DOI: https://doi.org/10.1242/jcs.s2-28.112.431

Trewick, S. (1998). Sympatric cryptic species in New Zealand Onychophora. Biological Journal of the Linnean Society, 63(3), 307-329. DOI: https://doi.org/10.1111/j.1095-8312.1998.tb01520.x

Trewick, S.A. (1999). Molecular diversity of Dunedin Peripatus (Onychophora: Peripatopsidae). New Zealand Journal of Zoology, 26(4), 381-393. DOI: https://doi.org/10.1080/03014223.1999.9518201

Trewick, S.A. (2000). Mitochondrial DNA sequences support allozyme evidence for cryptic radiation of New Zealand Peripatoides (Onychophora). Molecular Ecology, 9(3), 269-281. DOI: https://doi.org/10.1046/j.1365-294x.2000.00873.x

Vaidya, G., Lohman, D.J., & Meier, R. (2011). SequenceMatrix: concatenation software for the fast assembly of multigene datasets with character set and codon information. Cladistics, 27(2), 171-180. DOI: https://doi.org/10.1111/j.1096-0031.2010.00329.x

Zilch, A. (1954). Onychophoren aus El Salvador. Senckenbergiana Biologica, 35, 147-150.

Zitani, N.M., Thorn, R.G., Hoyle, M., Schulz, J.M., Steipe, T., Ruiz, Y.B., Sarquis-Adamson, Y., & Wishart, A.E. (2018). An Onychophoran and Its Putative Lepidopteran Mimic in the Arboreal Bryosphere of an Ecuadorian Cloud Forest. American Entomologist, 64(2), 94-101. DOI: https://doi.org/10.1093/ae/tmy025

Published

2020-03-31