Abstract
Our ability to reconstruct the evolutionary history of deep-water scalpellid barnacles (Crustacea: Cirripedia: Scalpellidae) and to understand their geographical distributions and ecology, is limited by the lack of a robust classification system. Analysis of the ontogeny of scalpellids highlights the significant morphological changes that occur during development from the smallest, recently settled juveniles up to full-grown specimens. Externally, these ontogenetic alterations can include changes in size, shape, and in the capitulum/peduncle ratio, as well as changes in the shape of plates resulting from modifications in the patterns of calcification. Using an exemplar developmental series of Mesoscalpellum carinatum (Hoek, 1881), we reveal how the misinterpretation of such changes has resulted in the recognition of polyphyletic and paraphyletic taxonomic units and how this has impacted all previous classifications proposed for the Scalpellidae. Our study explores a novel approach to the recognition of Mesoscalpellum Hoek, 1907 sensu Newman & Ross, 1971 as a monophyletic unit, based on the incorporation of data derived from developmental patterns. The re-establishment of this genus leads to significant systematic rearrangement. Multiple taxa currently placed in three scalpellid subfamilies and five genera were recognised as synonyms of just three Mesoscalpellum species—M. carinatum (Hoek, 1881), Mesoscalpellum sanctaebarbarae (Pilsbry, 1907) and Mesoscalpellum japonicum (Hoek, 1883). Two scalpellid genera, Barbascalpellum Zevina, 1978 and Pteroscalpellum Zevina, 1978, are no longer regarded as valid. In future, if study of scalpellid developmental series demonstrates that the holotype of a species is ambiguous and cannot be used for the precise application of the name of a taxon, a level of type status should be assigned to the full-grown specimen to promote its use for comparative study. We suggest that our perspective will help to resolve the confusion around the traditional typological grouping of other scalpellid genera and will foster the creation of a robust scalpellid classification.
References
- Anderson, D.T. (1994) Barnacles: structure, function, development and evolution. Chapman & Hall, London, Glasgow, New York, Tokyo, Melbourne and Madras, 357 pp.
- Annandale, N. (1906) Natural history notes from the R.I.M.S. ‘‘Investigator’’, Captain I.H. Heming, R.N., commanding. Series III, No 12. Preliminary report on the Indian Stalked Barnacles. The Annals and Magazine of Natural History, Series 7, 17, 389–400. https://doi.org/10.1080/00222930608562544
- Annandale, N. (1910) Description of a new species of Scalpellum from the Andaman Sea. Records of the Indian Museum, 5 (2), 115–116. https://doi.org/10.26515/rzsi/v5/i2/1910/163752
- Annandale, N. (1913) The Indian Barnacles of the subgenus Scalpellum. Records of the Indian Museum, 9 (4), 227–236. https://doi.org/10.26515/rzsi/v9/i4/1913/163654
- Aurivillius, C.W.S. (1898) Cirrhipedes nouveaux provenant des campagnes scientifiques de S.A.S. le Prince de Monaco. Bulletin de la Societe Zoologique de France, 23, 189–198.
- Barnard, K.H. (1924) Contributions to the Crustacean Fauna of South Africa. Cirripedia. Annals of the South African Museum, 20 (1), 1–103.
- Broch, H. (1953) Cirripedia Thoracica. The Danish Ingolf-Expedition, 3, 1–17.
- Calman, W.T. (1918) On barnacles of the genus Scalpellum from deep-sea telegraph cables. Annals and Magazine of Natural History, Series 9, 1 (1), 96–124. https://doi.org/10.1080/00222931808562293
- Chan, B.K.K., Prabowo, R.E. & Lee, K.-S. (2009) Crustacean fauna of Taiwan: barnacles. Vol. I. Cirripedia: Thoracica excluding the Pyrgomatidae and Acastinae. National Taiwan Ocean University, Keelung, 297 pp.
- Chan, B.K.K., Prabowom, R.E. & Lee, K.S. (2010) North West Pacific deep-sea barnacles (Cirripedia, Thoracica) collected by the TAIWAN expeditions, with descriptions of two new species. Zootaxa, 2405 (1), 1–47. https://doi.org/10.11646/zootaxa.2405.1.1
- Chan, B.K.K., Corbari, L., Rodriguez Moreno, P. & Jones, D. (2014) Two new deep-sea stalked barnacles, Arcoscalpellum epeeum sp. nov. and Gymnoscalpellum indopacificum sp. nov., from the Coral Sea, with descriptions of the penis in Gymnoscalpellum dwarf males. Zootaxa, 3866 (2), 261–276. https://doi.org/10.11646/zootaxa.3866.2.5
- Chan, B.K.K. & Rodriguez Moreno, P.A. (2020) A new deep-see barnacle, Calantica australpacifica sp. nov. (Scapelliformes: Calanticidae) in the South Pacific: first report on the presence of dwarf males on the sub-rostral position of Calantica. In: Laure Corbari, L., Shane, T., Ahyong, S.T & Tin-Yam Chan, T.Y. (Eds.), Tropical Deep-Sea Benthos. Vol. 31. Deep-sea crustaceans from Papua New Guinea. Muséum national d’Histoire naturelle, Paris, pp. 447–460.
- Chan, B.K.K, Dreyer, N., Gale, A.S., Glenner, H., Ewers-Saucedo, C., Pérez-Losada, M., Kolbasov, G.A., Crandall, K.A. & Høeg, J.T. (2021) The evolutionary diversity of barnacles, with an updated classification of fossil and living forms. Zoological Journal of the Linnean Society, 193 (3), 789–846. https://doi.org/10.1093/zoolinnean/zlaa160
- Darwin, C. (1851) A monograph of the subclass Cirripedia, with figures of all the species. The Lepadidae; or, pedunculated cirripedes. Ray Society, London, 400 pp. [published in 1852] https://doi.org/10.1080/03745485709495075
- Dreyer, N., Yusa, Y., Gale, A., Melzer, R. R., Yamato, S. & Hoeg, J. T. (2018) In the footsteps of Darwin: dwarf male attachment sites in scalpellid barnacles (Crustacea: Cirripedia: Thoracica) – implications for phylogeny and the evolution of sexual systems. Zoological Journal of the Linnean Society, 184, 999–1023. https://doi.org/10.1093/zoolinnean/zly018
- Foster, B.A. (1978) The Marine Fauna of New Zealand: Barnacles (Cirripedia: Thoracica). New Zealand Oceanographic Institute Memoir, 69, 1–160.
- Foster, B.A. (1980) Further records and classification of scalpellid barnacles (Cirripedia: Thoracica) from New Zealand. New Zealand Journal of Zoology, 7, 523–531. https://doi.org/10.1080/03014223.1980.11760684
- Foster, B.A & Buckeridge, J.S. (1995) Barnacles (Cirripedia: Thoracica) of seas off the Straits of Gibraltar. Bulletin du Museum National d’Histoire Naturelle. Section A: Zoologie Biologie et Ecologie Animales, 17, 163–191.
- Gale, A.S. (2016) Phylogeny of the deep-sea cirripede family Scalpellidae (Crustacea, Thoracica) based on shell capitular plate morphology. Zoological Journal of the Linnean Society, 176, 266–304. https://doi.org/10.1111/zoj.12321
- Gruvel, A. (1920) Cirrhipèdes provenant des campagnes scientifiques de S.A.S. le Prince de Monaco, 1885–1931. Résultats des Campagnes Scientifiques accomplies sur son Yacht par Albert 1er Prince de Monaco, 53, 1–89.
- Hendrickx, M.E. & Innocenti, G. (2019) Goose barnacles (Cirripedia: Thoracica: Lepadiformes and Scalpelliformes) from western Mexico. Cahiers de Biologie Marine, 60, 59–68. https://doi.org/10.21411/CBM.A.D2E0DE7D
- Hiro, F. (1933) Report on the Cirripedia collected by the surveying ships of the Imperial Fisheries Experimental Station on the continental shelf bordering Japan. Records of the Oceanographic Works in Japan, 5, 11–84.
- Hoek, P.P.C. (1883) Report on the Cirripedia collected by H.M.S. “Challenger” during the years 1873–1876. Systematic part. Report on the Scientific Results of the Voyage H.M.S. “Challenger”, during the years 1873–1876, Zoology, Part 25 (8), 1–169.
- Hoek, P.P.C. (1907) The Cirripedia of the Siboga expedition A. Cirripedia Pedunculata. Siboga Expeditie Monographe, 31 (a), 1–127. https://doi.org/10.5962/bhl.title.12959
- Leach, W.E. (1817) Distribution, systématique de la class Cirripèdes: par la même. Journal de Physique de Chimie et d’Histoire naturelle, 85, 67–69
- Lin, H., Høeg, J.T., Yusa, Y. & Chan, B.K.K. (2015) The origins and evolution of dwarf males and habitat use in thoracican barnacles. Molecular Phylogenetics and Evolution, 91, 1–11. https://doi.org/10.1016/j.ympev.2015.04.026
- Linnaeus, C. (1767). Systema naturae per regna tria naturae: secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Vol. 1. 12th Edition. Regnum Animale. 1 & 2. Laurentii Salvii, Holmiae [Stockholm], 532 pp.
- MacDonald, R. (1929) A report on some cirripedes collected by S.S. “Albatross” in the Eastern Pacific during 1892 and 1904. Bulletin of the Museum of Comparative Zoology at Harvard College, 69 (15), 527–538.
- Newman, W.A. (1987) Evolution of Cirripedes and their major groups. In: Southward, A.J. (Ed.), Crustacean Issues. Vol. 5. Barnacle Biology. Balkema, Rotterdam, pp. 3–42. https://doi.org/10.1201/9781315138053-1
- Newman, W.A. & Ross A. (1971) Antarctic Cirripedia. Antarctic Research Series 14. American Geophysical Union, Washington, 257 pp. https://doi.org/10.1029/ar014
- Newman, W.A. & Ross, A. (1998). Peduncular armament in the Scalpellomorpha (Cirripedia) and a new abyssal species from the East Pacific Rise. Journal of Crustacean Biology, 18 (3), 572–580. https://doi.org/10.1163/193724098x00395
- Pilsbry, H.A. (1890) Description of a new Japanese Scalpellum. Proceedings of the Academy of Natural Sciences of Philadelphia, 42, 441–443.
- Pilsbry, H.A. (1907) The Barnacles (Cirripedia) contained in the collections of the U. S. National Museum. Bulletin of the United States National Museum, 60, 1–122. https://doi.org/10.5479/si.03629236.60.1
- Pilsbry, H.A. (1908) On the classification of scalpelliform barnacles. Proceedings of the Academy of Natural Sciences of Philadelphia, 60, 104–111.
- Pilsbry, H.A. (1911) Barnacles of Japan and Bering Sea. Bulletin of the United States Bureau of Fisheries, 29, 59–84.
- Pilsbry, H.A. (1916) The sessile barnacles (Cirripedia) contained in the collections of the U.S. National Museum; including a monograph of the American species. Bulletin of the United States National Museum, 93, 1–366. https://doi.org/10.5479/si.03629236.93.1
- Pitriana, P., Jones, Diana S., Corbari, L., von Rintelen, K. (2020) New insights gained from museum collections: Deep-sea barnacles (Crustacea, Cirripedia, Thoracica) in the Museum National d’Histoire Naturelle, Paris, collected during the Karubar expedition in 1991. Zoosystematics and Evolution, 96 (2), 649–698. https://doi.org/10.3897/zse.96.55733
- Poltarukha, O.P. (2013) Barnacles (Cirripedia: Thoracica) from the Equatorial East Pacific. Russian Journal of Marine Biology, 39 (1), 51–57. [in Russian] https://doi.org/10.1134/s106307401301007
- Rao, M.V.L. & Newman, W.A. (1972) Thoracica Cirripedia from guyots of the mid-Pacific mountains. Transactions of the San Diego Society of Natural History, 17 (6), 69–94. https://doi.org/10.5962/bhl.part.19960
- Ren, X. (1989) On a collection of Cirripedia Thoracica from Madagascar and adjacent waters. Bulletin du Muséum national d’Histoire naturelle, Paris, Séries A: Zoologie, Biologie et Ecologie Animales, 11, 431–468.
- Ren, X. & Sha, Z. (2014) Description of a new genus and species of the subfamily Arcoscalpellinae Zevina, 1978 (Cirripedia: Thoracica: Scalpellidae) from deep waters of the South China Sea. Journal of Natural History, 48, 1055–1060. https://doi.org/10.1080/00222933.2013.841300
- Sander, K. (1983) The evolution of patterning mechanisms: gleanings from insect embryogenesis and spermatogenesis. In: Goodwin, B.C., Holder, N. & Wylie, C.C. (Eds.), Evolution and development. Cambridge University Press, Cambridge, pp. 137–159.
- Schrödl, M. & Haszprunar, G. (2016) Do we need Epitypes in Zoology? Spixiana, 39 (2), 199–201.
- Seguenza, G., (1873–1876) Ricerche paleontologiche intorno ai Cirripedi terziarii della provincia di Messina. Con appendice intorno ai Cirripedi viventi nel Mediterraneo, e sui fossili terziarii dell’Italia meridionale. Parte 1. Balanidi e Verrucidi; Parte 2. Terza famiglia, Lepadidi. Atti del’Accademia pontaniana, 10, 265–481.
- Shalaeva, K. (1996) The formation of external cuticular coat and plates in metamorphosis of barnacles (Crustacea, Thoracica) cypris larvae. Zoologicheskii Zhurnal, 75 (7), 995–1006. [in Russian]
- Shalaeva, K. & Boxshall, G. (2014) An illustrated catalogue of the scalpellid barnacles (Crustacea: Cirripedia: Scalpellidae) collected during the HMS “Challenger” expedition and deposited in the Natural History Museum, London. Zootaxa, 3804 (1), 1–63. https://doi.org/10.11646/zootaxa.3804.1.1
- Shalaeva, K. & Newman, W.A. (2016) Zevinaella—a new barnacle genus (Scalpellomorpha: Arcoscalpellinae) associated with crinoids (Echinodermata) from the Caribbean. Zootaxa, 4072(2), 151–170. https://doi.org/10.11646/zootaxa.4072.2
- Stubbings, H.G. (1936) Cirripedia. The John Murray Expedition 1933-34. Scientific Reports, Zoology, 4 (1), 1–70.
- Stubbings, H.G. (1961) Cirripedia Thoracica from tropical West Africa. Scientific results of the Danish Expedition to the coasts of tropical West Africa, 1945–1946, Atlantide Reports, 6, 7–41.
- Tarasov, N.E. & Zevina, G.B. (1957) Cirripedia Thoracica of the Seas of the USSR. Fauna SSSR 69. Zoologicheskii Institut Akademii Nauk SSSR, Leningrad, 268 pp. [in Russian]
- Turland, N.J., Wiersema, J.H., Barrie, F.R., Greuter, W., Hawksworth, D.L., Herendeen, P.S., Knapp, S., Kusber, W.-H., Li, D.-Z., Marhold, K., May, T.W., McNeill, J., Monro, A.M., Prado, J., Price, M.J. & Smith, G.F. (Eds.) (2018) International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Vegetabile, 159. Koeltz Botanical Books, Glashütten. Avaialble from: https://www.iapt-taxon.org/nomen/main.php (accessed 8 July 2024) https://doi.org/10.12705/Code.2018
- Weisbord, N.E. (1977) Scalpellid barnacles (Cirriredia) of Florida and surrounding waters. Bulletin of American Paleontology, 72 (299), 235–311.
- Withers, T.H. (1935) Catalogue of fossil Cirripedia in the Department of Geology. Vol. 2. Cretaceous. British Museum (Natural History), London, 534 pp.
- Withers, T.H. (1953) Catalogue of fossil Cirripedia in the Department of Geology. Vol. 3. Tertiary. British Museum (Natural History), London, 396 pp.
- Young, P.S. (1998) The Cirripedia (Crustacea) collected by the “Fisheries Steamer Meteor” in the eastern Atlantic. Arquivos do Museu Nacional Rio de Janeiro, 58, 1–5.
- Young, P.S. (2001) Redescription of Scalpellopsis striatociliata Broch, 1922, with a discussion on its phylogenetic position and the peduncular plate pattern in Scalpellids. Journal of Crustacean Biology, 21 (2), 456–468. https://doi.org/10.1163/20021975-99990147
- Young, P.S. (2007) The Scalpellomorpha (Crustacea, Cirripedia), with a list of extant species (except the Calanticidae). Galathea Report, 21, 7–73.
- Zevina, G.B. (1970) Cirripedia Thoracica from North West Pacific. Trudy Instituta Oceanologii Akademii Nauk SSSR, 86, 252–276. [in Russian]
- Zevina, G.B. (1975) Cirripedia Thoracica of the American Western Central Atlantic. Trudy Instituta Oceanologii Akademii Nauk SSSR, 100, 233–258. [in Russian]
- Zevina, G.B. (1978a) A new classification of the family Scalpellidae Pilsbry (Cirripedia, Thoracica). Part 1. Subfamilies Lithotryinae, Calanticinae, Pollicipinae, Scalpellinae, Brochiinae and Scalpellopsinae. Zoologicheskii Zhurnal, 57 (7), 998–1006. [in Russian]
- Zevina, G.B. (1978b) A new classification of the Scalpellidae (Cirripedia, Thoracica). Part 2 Subfamilies Arcoscalpellinae and Meroscalpellinae. Zoologicheskii Zhurnal, 57 (9), 1343–1352. [in Russian]
- Zevina, G.B. (1981) Cirriped crustaceans of the suborder Lepadomorpha (Cirripedia, Thoracica) of the World Ocean. Part 1. Family Scalpellidae. Opredeliteli po Faune SSSR, Leningrad, 398 pp. [in Russian]
- Zevina, G.B. (1982) Cirriped crustaceans of the suborder Lepadomorpha (Cirripedia, Thoracica) of the World Ocean. Part 2. Opredeliteli po Faune SSSR, Leningrad, 223 pp. [in Russian]