Skip to main content Skip to main navigation menu Skip to site footer
Responsive image
Issue: Vol. 5679 No. 2: 13 Aug. 2025
Type: Article
Published: 2025-08-13
DOI: 10.11646/zootaxa.5679.2.3
Page range: 217-242
Abstract views: 65
PDF downloaded: 1

Two new species of Clavellotis (Copepoda: Lernaeopodidae) with an approach to the phylogeny of the genus and its relationships inside the Clavella-Branch

Departamento de Ciencias Naturales y Ciencias Ambientales; Facultad de Ciencias del Mar; Universidad de Antofagasta; Casilla 170; Antofagasta; Chile.
Centro de Estudios Parasitológicos y Vectores (CEPAVE); Consejo Nacional de Investigaciones Cientificas y Tecnicas; Universidad Nacional de la Plata; Comision de Investigaciones Cientificas de la Provincia de Buenos Aires; calle 120 s/n e/60 y 64; CP 1900; La Plata; Buenos Aires; Argentina.
Ministry of Food; Agriculture and Livestock; Food Control Laboratory Directorate; Denizli; 20010; Turkey.
Centro de Estudios Parasitológicos y Vectores (CEPAVE); Consejo Nacional de Investigaciones Cientificas y Tecnicas; Universidad Nacional de la Plata; Comision de Investigaciones Cientificas de la Provincia de Buenos Aires; calle 120 s/n e/60 y 64; CP 1900; La Plata; Buenos Aires; Argentina.
Centro de Estudios Parasitológicos y Vectores (CEPAVE); Consejo Nacional de Investigaciones Cientificas y Tecnicas; Universidad Nacional de la Plata; Comision de Investigaciones Cientificas de la Provincia de Buenos Aires; calle 120 s/n e/60 y 64; CP 1900; La Plata; Buenos Aires; Argentina.
Departamento Ciencias Básicas. Facultad de Ciencias. Universidad Santo Tomas; Antofagasta; 399; Chile.
Crustacea Clavellotis Clavella branch relationships

Abstract

The identity of Clavellotis specimens parasitizing Chilean fishes has been determined using integrative taxonomy. Clavellotis dubius sp. nov., parasitic on Chromis crusma, can be distinguished from other species in the genus by a combination of characters, particularly the projection at the base of the cephalothorax, which shows variation related to the original aliform projection in Clavellotis. Additionally, its trunk is more elongated compared to the suborbicular trunk typical of other species. The male morphology aligns with the typical characteristics of the genus. The COI mtDNA genetic divergence between C. dubius sp. nov. and other Clavellotis ranges from 11.54% to 14.57%. Clavellotis girellae sp. nov., parasitic on Girella laevifrons, is morphologically similar to Clavellotis dilatata. However, males exhibit a unique dorsal projection shared only with Clavellotis sebastidis. The COI mtDNA genetic divergence for C. girellae sp. nov. ranges from 5.29% (with C. dilatata) to 12.94% (with C. sebastidis). With the addition of C. dubius and C. girellae, the number of Clavellotis species reported from Chilean fauna in the South Pacific increases to three, and the total number of species in the genus raises to 13. A phylogenetic analysis of Clavellotis is presented, including species from the South Pacific (C. dilatata, C. dubius sp. nov. and C. girellae sp. nov.), from South Atlantic (C. sebastidis) and Turkish Sea (Clavellotis fallax, Clavellotis strumosa, Clavellotis pagri and Clavellotis sargi). The analysis confirms the monophyly of Clavellotis, with C. sargi appearing as the most basal of the species within the genus. Additionally, the relationships of Clavellotis with other genera in the Clavella branch Alella, Clavella, Clavellisa Maxiclavella and Praeclavella—are also discussed.

 

References

  1. Adday, T. (2013) Parasitic crustacean of some marine fishes of Basrah Province, Iraq. Tesis de maestría, Agriculture College of University of Basrah, Basrah. [unknown pagination]
  2. Akmirza, A. (2000) Seasonal distribution of parasites detected in fish belonging to the Sparidae family found near Gökçeada. Turkish Journal of Parasitology, 24, 435–441.
  3. Alves, D.R. & Luque, J.L. (2001) Community ecology of the metazoan parasites of white croaker, Micropogonias furnieri (Osteichthyes: Sciaenidae), from the coastal zone of the State of Rio de Janeiro, Brazil. Memórias do Instituto Oswaldo Cruz, 96 (2), 145–153. https://doi.org/10.1590/S0074-02762001000200002
  4. Alves, D.R. & Luque, J.L. (2001) Aspectos quantitativos das infrapopulações de metazoários parasitos de Micropogonias furnieri (Osteichthyes: Sciaenidae) do litoral do Estado do Rio de Janeiro, Brasil. Parasitologia al Dia, 25, 30–35. https://doi.org/10.4067/S0716-07202001000100006
  5. Alves, D.R., Luque, J.L. & Paraguassú, A.R. (2002) Community ecology of the metazoan parasites of Pink Cusk-eel, Genypterus brasiliensis (Osteichthyes: Ophidiidae) from the coastal zone of the State of Rio de Janeiro, Brazil. Memórias do Instituto Oswaldo Cruz, 97, 683–689. https://doi.org/10.1590/S0074-02762002000500017
  6. Barnard, K.H. (1955) South African parasitic Copepoda. Annals of the South African Museum, 41 (5), 223–312.
  7. Ben Hassine, K., Essafi, K. & Raibaut, A. (1978) Les lernaeopodidés, copépodes parasites de Sparidae de Tunisie. Archives de l’Institut Pasteur de Tunis, 55 (4), 431–454.
  8. Benmansour, B. & Ben Hassine, O.K. (1998) Preliminary analysis of parasitic copepod species richness among coastal fishes of Tunisia. Italian Journal of Zoology, 65, 341–344. https://doi.org/10.1080/11250009809386844
  9. Benmansour, B., Ben-Hassine, O.K., Diebakate, D. & Raibaut, A. (2001) Sur deux espèces des copépodes Lernaeopodidae (Siphonostomatoida) parasite du marbré Lithognathus mormyrus (Linnaeus, 1758) (Pisces, Sparidae). Zoosystema, 23, 695–703.
  10. Bere, R. (1936) Parasitic copepods from Gulf of Mexico fish. The American Midland Naturalist, 17 (3), 571–625. https://doi.org/10.2307/2419936
  11. Boualleg, C., Serid, M., Kaouachi, N., Quiliquini, Y. & Bensouilah, M. (2010) Les Copépodes parasites des poissons téléostéens du littoral Est-algérien. Bulletin de l’Institut Scientifique, Rabat, section Sciences de la Vie, 32 (2), 65–72.
  12. Boxshall, G.A. & Halsey, S.H. (2004) An introduction to copepod diversity. Ray Society, London, 966 pp.
  13. Brian, A. (1898) Catalogo di copepodi parassiti dei pesci della Liguria. Bolletino dei musei e laboratorii di zoologia e di anatomia comparata della R. Università di Genova, 61, 1–27.
  14. Brian, A. (1899) Di alcuni Crostacei parassiti dei pesci dell'Isola d'Elba. Atti della Società Ligustica di Scienze Naturali e Geografiche, 10, 3–10.
  15. Brian, A. (1902) Note su alcuni Crostacei parassiti dei pesci del Mediterraneo. Atti della Società Ligustica di Scienze Naturali e Geografiche, 13, 30–45.
  16. Brian, A. (1903) Sui Copepodi parassiti di pesci marini dell’Isola d’Elba. 4è nota. Bolletino dei musei e laboratorii di zoologia e di anatomia comparata della R. Università di Genova, 5 (121), 1–8.
  17. Brian, A. (1905) Sur copepododes raccolti nel Golfo di Napoli da Oronzio G. ed Achille Costa. Annuario del Regio Museo Zoologico della Reale Università di Napoli, Nuova Serie, 1 (24), 1–11.
  18. Brian, A. (1906) Copepodi parassiti dei Pesci d'Italia. Stab. Tipo-Litografico R. Istituto Sordomuti, Turin. [unknown pagination] https://doi.org/10.5962/bhl.title.58642
  19. Brian, A. (1924) Copepoda. Copepodes commensaux et parasites des cotes Mauritaniennes. In: Parasitologica Mauritanica. Matériaux pour la faune parasitologique en Mauritanie, 1 (1), 1–66.
  20. Brian, A. (1924) Parasitologia mauritanica. Matériaux pour la faune parasitologique en Mauritanie. Arthropoda (1er partie). Copepoda. Bulletin du Comité d'Études Historiques et Scientifiques de l'Afrique Occidentale Française. Juill-sept, 1924, 365–427.
  21. Cabral, P. (1983) Morphologie, biologie et écologie des Copépodes parasites du Loup Dicentrarchus labrax (Linné, 1758) et du Sar rayé Diplodus sargus (Linné, 1758) de la région languedocienne. Tesis de 3ème cycle, Université Montpellier, Montpellier. [unknown pagination]
  22. Caillet, C. (1977) Contribution à l'étude des copépodes parasites de quelques poissons Téléostéens de la région de Sète. Rapport de stage D.E.A., Université Montpellier, Montpellier. [unknown pagination]
  23. Cantatore, D., Braicovich, P., Alarcos, A.J. & Timi, J.T. (2012) New records of parasitic copepods (Crustacea, Copepoda) from marine fishes in the Argentinean Sea. Acta Parasitologica, 57 (1), 83–89. https://doi.org/10.2478/s11686-012-0010-0
  24. Capart, A. (1959) Copépodes parasites. Résultats Scientifiques de l'Expédition Océanographique Belge dans les Eaux Côtières Africaines de l'Atlantique Sud- (1948–1949), 3, 55–126.
  25. Castro Romero, R. & Baeza Kuroki, H. (1984) Clavellotis, new genus (Copepoda: Lernaeopodidae), and redescription of Clavellotis dilatata (Kroyer, 1863). Journal of Crustacean Biology, 4 (4), 688–694. https://doi.org/10.2307/1548077
  26. Castro Romero, R. & González, M.T. (2005) Clavellotis sebastidis sp. nov. (Copepoda, Lernaeopodidae) parasitic on Sebastes oculatus Valenciennes, 1833 from Argentina. Acta Parasitologica, 50 (1), 74–79. https://doi.org/10.2478/s11686-005-0010-x
  27. Castro Romero, R., Montes, M.M. & Martorelli, S. (2022) Maxiclavella and Praeclavella (Siphonostomatoida: Lernaeopodidae) new genera confirmed by molecular and morphological evidence. Anais da Academia Brasileira de Ciências, 94 (4), 1–20. https://doi.org/10.1590/0001-3765202220200992
  28. Castro, R., Montes, M., Martorelli, S.R., Sepúlveda, D., Tapia, S. & Martinez-Aquino, A. (2016) Integrative taxonomy of Peniculus, Metapeniculus and Trifur (Siphonostomatoida: Pennellidae) copepod parasites of marine fishes from Chile: Species delimitation analysis using DNA barcoding evidence. Systematics and Biodiversity, 14 (5), 1–18. https://doi.org/10.1080/14772000.2016.1158213
  29. Dippenaar, S.M. (2016) Does Alella Leigh-Sharpe, 1925 (Siphonostomatoida: Lernaeopodidae) really consist of seven species? Systematic Parasitology, 93 (1), 47–56. https://doi.org/10.1007/s11230-015-9601-0
  30. Dippenaar, S.M. (2024) Twenty years later: Biodiversity of marine symbiotic Siphonostomatoida (Copepoda) off Southern Africa. Zootaxa, 5419 (1), 85–111. https://doi.org/10.11646/zootaxa.5419.1.3
  31. Do, T.T. & Ho, J.S. (1983) Clavellopsis nodula sp. nov. (Copepoda: Lernaeopodidae) parasitic on sea bream, Mylio macrocephalus (Basilewsky) (Pisces: Sparidae) in Japan. Fish Pathology (Gyobyo Kenkyu), University of Tokyo, 18 (1), 31–36. https://doi.org/10.3147/jsfp.18.31
  32. Erasmus, A., Hadfield, K.A., Wepener, V. & Smit, N.J. (2023) Morphological and molecular characterization of Alella igillimpethu sp. nov. (Copepoda: Siphonostomatoida: Lernaeopodidae) parasitising the southern African endemic intertidal klipfish, Clinus superciliosus. Systematic Parasitology, 100 (1), 69–83. https://doi.org/10.1007/s11230-022-10071-3
  33. Folmer, O., Black, M., Hoeh, W., Lutz, R. & Vrijenhoek, R. (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3 (5), 294–299.
  34. Gasmi, Y., Belhocine, K., Abdeli, R. & Khati, W. (2017) Parasitic specificity in the sparidae family fish Pagellus erythrinus of the El Kala coast, Algeria. AACL Bioflux, 10 (4), 721–738.
  35. González, M.T. & Moreno, C.A. (2005) The distribution of the ectoparasite fauna of Sebastes capensis from the southern hemisphere does not correspond with zoogeographical provinces of free-living marine animals. Journal of Biogeography, 32, 1539–1547. https://doi.org/10.1111/j.1365-2699.2005.01323.x
  36. Heller, C. (1865) Crustaceen. [Crustacea]. Reise der Österreichischen Fregatte Novara um die Erde, in den Jahren 1857, 1858, 1859 unter den Befehlen des Commodore B. von Wüllerstorf-Urbair, Zoologischer Theil, Kaiserlich-Königliche Hof- und Staatsdruckerei, Wien, Band. 2 (Abtheilung 3), 1–280.
  37. Hewitt, G.C. & Blackwell, R.G. (1987) A new species of Alella (Copepoda, Lernaeopodidae) parasitic in the tarakihi, Cheilodactylus macropterus, in New Zealand. New Zealand Journal of Marine and Freshwater Research, 21 (1), 141–147. https://doi.org/10.1080/00288330.1987.9516208
  38. Iannacone, J. (2004) Metazoos parásitos de la mojarrilla Stellifer minor (Tschudi) (Osteichthyes, Sciaenidae) capturados por pesquería artesanal en Chorrillos, Lima, Perú. Revista Brasileira de Zoologia, 21, 815–820. https://doi.org/10.1590/S0101-81752004000400015
  39. Kabata, Z. (1979) Parasitic Copepoda of British Fishes. The Ray Society, London, 468 pp.
  40. Kabata, Z. (1990) Revision of the genus Clavellopsis Wilson, 1915 (Copepoda: Lernaeopodidae). Canadian Journal of Zoology, 68, 2564–2566. https://doi.org/10.1139/z90-357
  41. Katoh, K., Rozewicki, J. & Yamada, K.D. (2019) MAFFT online service: Multiple sequence alignment, interactive sequence choice and visualization. Briefings in Bioinformatics, 20, 1160–1160. https://doi.org/10.1093/bib/bbx108
  42. Kensley, B. & Grindley, J.R. (1973) South African Parasitic Copepoda. The Annals of the South African Museum, 62 (3), 69–130.
  43. Koyuncu, C., Romero, R. & Genç, E. (2014) Clavellotis briani (Copepoda, Lernaeopodidae) Infestation on Striped Seabream, Lithognathus mormyrus (Sparidae) from the Northeast Mediterranean Sea, Turkey. Journal of Agricultural Sciences, 21 (1), 152–157. https://doi.org/10.15832/tbd.71968
  44. Krøyer, H. (1863–1864) Bidrag til Kundskab om Snyltekrebsene. Naturhistorisk Tidsskrift Ser. III, 2 (1/2 & 3), 75–320, pls. 1–9 [1863] & 321–426. https://doi.org/10.5962/bhl.title.59539
  45. Kumar, S., Stecher, G., Li, M., Knyaz, C. & Tamura, K. (2018) MEGA X: Molecular Evolutionary Genetics Analysis across Computing Platforms. Molecular Biology and Evolution, 35, 1547–1549. https://doi.org/10.1093/molbev/msy096
  46. Kurz, W. (1877) Studien über die Familie der Lernaepodiden. Zeitschrift für Wissenschaftliche Zoologie, 29, 380–426.
  47. Lanfear, R., Calcott, B., Kainer, D., Mayer, C. & Stamatakis, A. (2014) Selecting optimal partitioning schemes for phylogenomic datasets. BMC Evolutionary Biology, 14, 1–82. https://doi.org/10.1186/1471-2148-14-82
  48. Larrañeta, M.G. (1964) Sobre la biología de Pagellus erythrinus (L.) especialmente del de las costas de Castellón. [On the biology of Pagellus erythrinus (L.), especially that of the Castellón coast]. Investigación Pesquera, Barcelona, 27, 131–146.
  49. Luque, J.L. (1994) Dinámica poblacional y estructura de la comunidad de metazoarios parásitos de Menticirrhus ophicephalus (Pisces: Sciaenidae) en la costa peruana. Revista de Biología Tropical, Costa-Rica, 42, 21–29.
  50. Luque, J.L. (1996) Copépodos parásitos do Pargo, Pagrus pagrus (Linnaeus) (Osteichthyes, Sparidae) do litoral do estado do Rio de Janeiro, Brasil. Resumos XXI Congreso Brasileiro de Zoología, 1996, 50.
  51. Luque, J.L. & Takemoto, R.M. (1996) Parasitic Copepods on Orthoptists ruber (Cuvier) and Haemulon steindachneri (Jordan and Gilbert) (Osteichthyes: Haemulidae) from the Brazilian coast, with the description of a new species of Caligus (Siphonostomatoida: Caligidae). Revista Brasileira de Biologia, 56 (2), 529–546.
  52. Luque, J.L., Amato, J.F.R. & Takemoto, R.M. (1996) Comparative analysis of the communities of metazoan parasites of Orthopristis ruber and Haemulon steindachneri (Osteichthyes: Haemulidae) from the southeastern Brazilian littoral: I. Structure and influence of the size and sex of hosts. Revista Brasileira de Biologia, 56, 279–292.
  53. Luque, J.L., Cruces, C., Chero, J., Paschoal, F., Alves, P.V., Da Silva, A.C., Sanchez, L. & Iannacone, J. (2016) Checklist of metazoan parasites of fishes from Peru. Neotropical Helminthology, 10, 301–375. https://doi.org/10.24039/rnh2016102751
  54. Luque, J.L. & Farfán, C.A. (1990) Copépodos de la familia Lernaeopodidae (Siphonostomatoida) parásitos de algunos peces de la familia Sciaenidae (Teleostei) en el mar peruano. Parasitologia al Dia, 14 (3–4), 63–66.
  55. Lv, H.J. & Huang, Y. (2012) [Phylogenetic relationship among some groups of orthopterans based on complete sequences of the mitochondrial COI gene]. Dongwuxue Yanjiu, 3, 319–328. https://doi.org/10.3724/SP.J.1141.2012.03319
  56. Monod, T. (1923) Notes carcinologiques. (Parasites et commensaux). Bulletin de l'Institut Océanographique de Monaco, 427, 19–22.
  57. Nagasawa, K. (2011) A checklist of the parasitic copepods (Crustacea) of fishes and invertebrates of the Seto Inland Sea, Japan (1935–2011), with a new locality record of Caligus macarovi (Caligidae). Bulletin of the Hiroshima University Museum, 3, 113–128.
  58. Nei, M. & Kumar, S. (2000) Molecular evolution and phylogenetics. Oxford University Press. Oxford, 333 pp. https://doi.org/10.1093/oso/9780195135848.001.0001
  59. Nuñes-Ruivo, L. (1953) Copépodes parasites de poissons. [Copepods parasitic on fish.]. In: Travaux du C.L.O.E.C. d'Algérie No. 15. Campagne du "Professeur Lacaze-Duthiers" sur les côtes d'Algérie, (Juin-Juillet 1952). Vie et Milieu Supplément, 3, pp. 115–138.
  60. Öktener, A., Alaş, A. & Solak, K. (2008) Clavellotis strumosa (Brian, 1906) (Copepoda; Lernaeopodidae) a Gill Parasite of Pagellus erythrinus (Linnaeus, 1758) from the Sea of Marmara. Crustaceana, 81, 631–636. https://doi.org/10.1163/156854008784092193
  61. Öktener, A., Alaş, A. & Türker, D. (2017) Confirmed record of Clavellotis fallax (Heller) (Siphonostomatoida; Lernaeopodidae) from Dentex dentex (Linnaeus) with morphological characters in Turkey. Bonorowo Wetlands, 7 (1), 4–7. https://doi.org/10.13057/bonorowo/w070102
  62. Oliva, M.E. & Luque, J. (1998) Metazoan parasite infracommunities in five sciaenids from the central Peruvian Coast. Memórias do Instituto Oswaldo Cruz, 93 (2), 175–180. https://doi.org/10.1590/S0074-02761998000200007
  63. Oliva, M.E., Luque, J.L. & Iannacone, J.A. (1990) The Metazoan parasites of Stellifer minor (Tschudi, 1844): An Ecological Approach. Memórias do Instituto Oswaldo Cruz, 85 (3), 271–274. https://doi.org/10.1590/S0074-02761990000300003
  64. Papoutsoglou, S.E. (1976) Metazoan parasites of fishes from Saronicos Gulf Athens – Greece. Thalassographica, 1 (1), 69–102.
  65. Paraguassú, A.R., Luque, J.L. & Alves, D.R. (2002) Community ecology of the metazoan parasites of red porgy Pagrus pagrus (L. 1758) (Osteichthyes, Sparidae) from the coastal zone, state of Rio de Janeiro, Brazil. Acta Scientiarum, 24 (2), 461–467.
  66. Pearse, A.S. (1952) Parasitic Crustacea from the Texas coast. Publications of the Institute of Marine Science, Port Aransas, 1952, 1–42.
  67. Pillai, N.K. (1962) Copepods parasitic on South Indian fishes: Families Lernaeopodidae and Naobranchidae. Journal of the Marine Biological Association of India, 4, 58–94.
  68. Radujković, B.M. & Raibaut, A. (1989) Parasites des poissons marins des côtes du Monténégro: Copépodes. Acta Adriatica, 28, 237–278.
  69. Raibaut, A., Ben Hassine, O.K. & Maamouri, K. (1971) Copepodes Parasites des Poissons de Tunisie (premiere Serie). Bulletin de l'Institut National Scientifique et Technique d'Oceanographie et de Peche de Salammbo, 2 (2), 169–197.
  70. Ramdane, Z. & Trilles, J.P. (2007) Parasitic copepods (Crustacea: Copepoda) from Algerian marine fishes. Zootaxa, 1574 (1), 49–68. https://doi.org/10.11646/zootaxa.1574.1.3
  71. Ramos Alves, D. & Luque, J.L. (2000) Metazoários parasitos de Micropogonias furnieri (Osteichthyes: Sciaenidae) do litoral do Estado do Rio de Janeiro, Brasil. Parasitología al día, 24 (1–2), 40–45. https://doi.org/10.4067/S0716-07202000000100006
  72. Raupach, M.J., Barco, A., Steinke, D., Beermann, J., Laakmann, S., Mohrbeck, I., Neumann, H., Kihara, T.C., Pointner, K., Radulovici, A., Segelken-Voigt, A., Wesse, C. & Knebelsberger, T. (2015) The application of DNA barcodes for the identification of marine crustaceans from the North Sea and adjacent regions. PLoS ONE, 10 (9), e0139421. https://doi.org/10.1371/journal.pone.0139421
  73. Richiardi, S. (1880) Catalogo sistematico dei crostacei che vivono sul corpo degli animali aquatici. Catalogo Sez. Italiana Exposizione Internazionale di Pesca in Berlino, 147–152.
  74. Rodríguez-Santiago, M.A., Gómez, S., Rosales-Casián, J.A. & Grano-Maldonado, M.I. (2015) Copépodos parásitos del pez vermillon Sebastes miniatus (Pisces: Scorpaenidae) en las costas de Baja California (Pacífico Noreste). Neotropical Helminthology, 9 (1), 1–12. https://doi.org/10.24039/rnh201591774
  75. Rodríguez-Santiago, M.A., Rosales-Casián, J.A. & Grano-Maldonado, M.I. (2014) Dynamics of a parasite assemblage of the vermilion rockfish Sebastes miniatus from northwestern Baja California, Mexico. Helgoland Marine Research, 68 (2), 299–306. https://doi.org/10.1007/s10152-014-0389-0
  76. Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & 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. https://doi.org/10.1093/sysbio/sys029
  77. Roubal, F.R., Armitage, J. & Rohde, K. (1983) Taxonomy of metazoan ectoparasites of snapper, Chrysophrys auratus (family Sparidae), from southern Australia, eastern Australia and New Zealand. Australian Journal of Zoology, Supplemental Series, 94, 1–68. https://doi.org/10.1071/AJZS094
  78. Schwarz, G. (1978) Estimating the dimension of a model. Annals of Statistics, 6 (2), 461–464. https://doi.org/10.1214/aos/1176344136
  79. Sebone, M. & Dippenaar, S. (2024) Reports of Pennellidae Burmeister, 1835 (Copepoda: Siphonostomatoida) post-metamorphosed females off the coast of southern Africa. Systematic Parasitology, 101 (32), 1–20. https://doi.org/10.1007/s11230-024-10157-0
  80. Shiino, S. (1959) Sammlung der parasitischen Copepoden in der Präfekturuniversität von Mie. Reports of the Faculty of Fisheries, Mie University, 3, 334–374.
  81. Skinner, R.H. (1978) Some external parasites of Florida fishes. Bulletin of Marine Science, 28 (3), 590–595.
  82. Soares, I.A. & Luque, J.L. (2015) Seasonal variability of the composition and structure of parasites of real porgy Pagrus pagrus (Perciformes, Sparidae) off Brazil. Helmintologia, 52 (3), 236–243. https://doi.org/10.1515/helmin-2015-0038
  83. Soares, I.A., Lanfranchi, A.L., Luque, J.L., Haimovici, M. & Timi, J.T. (2018) Are different parasite guilds of Pagrus pagrus equally suitable sources of information on host zoogeography? Parasitology Research, 117, 1865–1875. https://doi.org/10.1007/s00436-018-5869-8
  84. Soares, I.A., Vieira, F.M. & Luque, J.L. (2014) Parasite community of Pagrus pagrus (Sparidae) from Rio de Janeiro, Brazil: Evidence of temporal stability. Revista Brasileira de Parasitologia Veterinária, 23, 216–223. https://doi.org/10.1590/S1984-29612014036
  85. Tkach, V.V., Littlewood, D.T.J., Olson, P.D., Kinsella, J.M. & Swiderski, Z. (2003) Molecular phylogenetic analysis of the Microphalloidea Ward, 1901 (Trematoda: Digenea). Systematic Parasitology, 56, 1–15. https://doi.org/10.1023/A:1025546001611
  86. Valle, A. (1880) Crostacei parassiti dei pesci del mare adriatico. Bolletino della Società Adriatica di Scienze Naturali in Trieste, 6, 55–90.
  87. Walter, T.C. & Boxshall, G. (2025) World of Copepods Database. World Register of Marine Species. Available from: https://www.marinespecies.org/aphia.php?p=taxdetails&id=135602 (accessed 15 February 2025)
  88. Wood, C.L., Micheli, F., Fernandez, M., Gelcich, S., Castilla, J.C. & Carvajal, J. (2013) Marine protected areas facilitate parasite populations among four fished host species of central Chile. Journal of Animal Ecology, 82, 1276–1287. https://doi.org/10.1111/1365-2656.12104
  89. Yamaguti, S. (1939) Parasitic copepods from fishes of Japan. Part 6. Lernaeopodoida, 1. Volume Jubilare pro Professore Sadao Yoshida, 2, 529–578.
  90. Youssef, F., Benmansour, B., Ben Hassine, O.K. & Tlig, S.Z. (2016) Some parasitic copepods of selected Teleost and Chondrichthyan fishes from the Tunisian gulfs. African Journal of Microbiology Research, 10 (35), 1467–1476. https://doi.org/10.5897/AJMR2015.7820

How to Cite

Romero, R.C., Montes, M.M., Ӧtkener A., Shimabukuro, M.I., Theiller, M. & Campos, L. (2025) Two new species of Clavellotis (Copepoda: Lernaeopodidae) with an approach to the phylogeny of the genus and its relationships inside the Clavella-Branch. Zootaxa, 5679 (2), 217–242. https://doi.org/10.11646/zootaxa.5679.2.3