Invertebrate Systematics Invertebrate Systematics Society
Systematics, phylogeny and biogeography

An overview of the phylogeny of the families Lafoeidae and Hebellidae (Hydrozoa : Leptothecata): their composition and classification

Antonio C. Marques A D , Alvaro L. Peña Cantero B and Alvaro E. Migotto C
+ Author Affiliations
- Author Affiliations

A Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, CP 11461, 05422-970, São Paulo, SP, Brazil.

B Instituto Cavanilles de Biodiversidad y Biología Evolutiva, Universidad de Valencia, Apdo. Correos 22085, 46071 Valencia, Spain.

C Centro de Biologia Marinha, Universidade de São Paulo, Caixa Postal 83, 11600-970 – São Sebastião, SP, Brazil.

D Corresponding author. Email:

Invertebrate Systematics 20(1) 43-58
Submitted: 6 July 2005  Accepted: 5 January 2006   Published: 17 March 2006


A cladistic analysis of the genera of the family ‘Lafoeidae’ was performed in order to investigate suprageneric classifications and the boundaries of the family, as well as to organise the available morphological data and discuss the possible evolution of some morphological characters. Our results suggest that the former ‘Lafoeidae’ must be separated into two families: the Hebellidae and the Lafoeidae (including the subfamilies Lafoeinae and Zygophylacinae).


The authors wish to thank Professor Wim Vervoort (Nationaal Natuurhistorisch Museum, Leiden, The Netherlands) for his help with the bibliography, Dr A. M. García-Carrascosa (Universidad de Valencia, Spain) for accommodating us in his laboratory for part of the studies, Dr Peter Schuchert (Museum d’Histoire Naturelle, Geneve, Switzerland) for his comments on a previous version of the manuscript, and to Dale R. Calder (Royal Ontario Museum, Canada) and an anonymous referee for their valuable suggestions. We also acknowledge the staff of the museums listed in Table 3 for the loan of species. The project had financial support from Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP 2001/02626-7, 2001/10677-0, 2003/02432-3, and 2004/09961-4), Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq 471960/2004-7), National Museum of Natural History, Department of Invertebrate Zoology, United States Antarctic Program funded by the National Science Foundation (OPP-9509761), from the Ministerio de Educación y Ciencia of Spain and the Fondo Europeo de Desarrollo Regional (FEDER) (CGL2004-04684/ANT), and from the National Science Foundation (EF-0531779). Antonio C. Marques and Alvaro E. Migotto also have financial support from the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq; 302596/2003-8 and 300194/1994-3 respectively); Álvaro L. Peña Cantero has a research contract funded by the Ministerio de Ciencia y Tecnología and the Universidad de Valencia, Spain (Ramón y Cajal Program).


Agassiz A. (1865). North American Acalephae. In ‘Illustrated Catalogue of the Museum of Camparative Zoölogie, at Harvard College, no. II’. (Cambridge University Press: Cambridge: MA, USA.)

Allman G. J. (1864) On the construction and limitation of genera among the Hydroida. Annals and Magazine of Natural History 13(77), 345–380.

Allman G. J. (1888) Report on the Hydroida dredged by H. M. S. Challenger during the years 1873–76. Part II – The Tubulariinae, Corymorphinae, Campanulariinae, Sertulariinae, and Thalamophora. Report on the Scientific Results of the Voyage of H. M. S. Challenger During the Years 1873–76. Zoology (Jena, Germany) 23, 1–90.

Andrade L. P., and Migotto A. E. (1997). Is there a link between Hebella hydroids (Hydrozoa, Lafoeidae) and Staurodiscus medusae (Hydrozoa, Laodiceidae)? Proceedings of the VII Colacmar – Congresso Latino-Americano sobre Ciências do Mar, Santos 35–36.

Antsulevich A. E. (1987). ‘Hydroids from the Shelf Waters of Kurile Islands’. (Zoologicheskii Institut, Akademiya Nauk: Leningrad, SSSR). [In Russian with English summary.]

Billard A. (1910) Revision d’une partie de la collection des hydroïdes du British Museum. Annales des Sciences Naturelles, Zoologie 11(9), 1–67.

Blanco O. M. (1968) Nueva contribución al conocimiento de la fauna marina hidroide. Revista del Museo de La Plata [Zoología] 10(87), 195–224.

Blanco O. M. (1976) Hidrozoos de la expedición Walther Herwig. Revista del Museo de La Plata [Zoología] 12(113), 27–74.

Boero F., and Bouillon J. (1989). An evolutionary interpretation of anomalous medusoid stages in the life cycles of some Leptomedusae (Cnidaria). In ‘Reproduction, Genetics and Distributions of Marine Organisms. Proceedings of the XXIII European Marine Biology Symposium’. (Eds J. S. Ryland and P. A. Tyler.) pp. 37–41.

Boero F., Bouillon J., Kubota K. (1997) The medusae of some species of Hebella Allman, 1888, and Anthohebella gen. nov. (Cnidaria, Hydrozoa, Lafoeidae), with a world synopsis of species. Zoologische Verhandelingen 310, 1–53.

Bouillon J. (1985) Essai de classification des Hydropolypes-Hydroméduses (Hydrozoa-Cnidaria). Indo-Malayan Zoology 2(1), 29–243.

Bouillon J. (1994). Classe des hydrozoaires (Hydrozoa Owen, 1843). In ‘Traité de Zoologie, Tome III, Fascicule 2 - Cnidaires et Ctenaires’. (Eds J. Bouillon, C. Carré, D. Carré, A. Franc, J. Goy, M. L. Hernandez-Nicaise, Y. Tiffon, D. van de Vyver and M. Wade.) pp. 29–416. (Masson: Paris, France.)

Calder D. R. (1991) Shallow-water hydroids of Bermuda: the Thecatae, exclusive of Plumularioidea. Life Science Contributions, Royal Ontario Museum 154, 1–140.

Calder D. R., Vervoort W. (1998) Some hydroids (Cnidaria: Hydrozoa) from the Mid-Atlantic Ridge, in the North Atlantic Ocean. Zoologische Verhandelingen 319, 1–65.

Carpenter J. M. (1988) Choosing among multiple equally parsimonious cladograms. Cladistics 4, 291–296.

Carpenter J. M. (1994) Successive weighting, reliability and evidence. Cladistics 10, 215–220.
CrossRef |

Clarke S. F. (1879) Report on the Hydroida collected during the exploration of the Gulf Stream and Gulf of Mexico by Alexander Agassiz, 1877–78. Bulletin of the Museum of Comparative Zoölogy at Harvard College 5(10), 239–252.

Cornelius P. F. S. (1975) A revision of the species of Lafoeidae and Haleciidae (Coelenterata: Hydroida) recorded from Britain and nearby seas. Bulletin of the British Museum (Natural History). Zoology 28, 375–426.

Cornelius P. F. S. (1995). ‘Synopses of the British Fauna (new series), no. 50. North-west European Thecate Hydroids and their Medusae.’ (Linnean Society of London: London, UK.)

Farris J. S. (1969) A successive approximations approach to character weightings. Systematic Zoology 18, 374–385.

Ferrarezzi H., and Marques A. C. (1997). Elementos Básicos de Sistemática Filogenética. In ‘Análise cladística numérica e recursos computacionais. 2a Edição’. (Ed. D. S. Amorim.) pp. 161–186. (Sociedade Brasileira de Entomologia/Holos Editora: Ribeirão Preto, Brasil.)

Fraser C. M. (1912) Some hydroids of Beaufort, North Carolina. Bulletin of the United States Bureau of Fisheries 30, 337–387.

Fraser C. M. (1937). ‘Hydroids of the Pacific coast of Canada and the United States.’ (The University of Toronto Press: Toronto, Canada.)

Fraser C. M. (1938) Hydroids of the 1932, 1933, 1935, and 1938 Allan Hancock Pacific Expeditions. Allan Hancock Pacific Expeditions 4(3), 129–153.

Fraser C. M. (1944). ‘Hydroids of the Atlantic coast of North America.’ (The University of Toronto Press: Toronto, Canada.)

Fraser C. M. (1946). ‘Distribution and Relationship in American Hydroids.’ (The University of Toronto Press: Toronto, Canada.)

Fraser C. M. (1948) Hydroids of the Allan Hancock Pacific Expeditions since March, 1938. Allan Hancock Pacific Expeditions 4(5), 179–343.

Hadži J. (1913) Poredbena hidroidska istraživanja. I. Hebella parasitica (Ciam.). (S. dodatkom: Hebellopsis brochi g. et sp. n. i Hebella (?) gigas Pieper). Rad Jugoslavenske Akademije Znanosti i Umjetnosti 198, 105–210.

Haeckel E. (1889) Symbiotic Hydrozoa living in the deep-sea Keratosa. Report on the Scientific Results of the Voyage of H. M. S. Challenger During the Years 1873–76. Zoology 32, 1–92.

Hartlaub C. (1905) Die Hydroiden der Magalhaensischen Region und chilenischen Küste.  Zoologische Jahrbücher 6(3), 497–714.

Hennig W. (1950). ‘Grundzüge einer Theorie der Phylogenestichen Systematik.’ (Deutscher Zentralverlag: Berlin, Germany.)

Hennig W. (1966). ‘Phylogenetic Systematics.’ (University of Illinois Press: Urbana, IL, USA.)

Hincks T. (1868). ‘A History of the British Hydroid Zoophytes. Volume 1 and 2.’ (John van Voorst: London, UK.)

Hirohito S. (1967). ‘A Review of the Hydroids of the Family Clathrozonidae with Description of a New Genus and Species from Japan.’ (Biological Laboratory, Imperial Household: Tokyo, Japan.)

Hirohito S. (1971). ‘Additional Notes on Clathrozoon wilsoni Spencer.’ (Biological Laboratory, Imperial Household: Tokyo, Japan.)

ICZN (1991). ‘International Code of Zoological Nomenclature.’ 4th edn. (The International Trust for Zoological Nomenclature: London, UK.)

Jäderholm E. (1903) Aussereuropäischen Hydroiden im schwedischen Reichsmuseum. Arkiv för Zoologi 1, 259–312.

Kitching I. L., Forey P. L., Humphries C. J., and Williams D. M. (1998). ‘Cladistics, A Practical Course in Systematics.’ 2nd edn. (The Systematics Association: London, UK.)

Kramp P. L. (1932) The Godthaab expedition 1928. Hydroids. Meddelelser om Grønland 79, 1–86.

Kühn A. (1913) Entwicklungsgeschichte und Verwandschaftsbeziehungen der Hydrozoen. I. Teil: Die Hydroiden. Ergebnisse und Fortschritte der Zoologie 4(1–2), 1–284.

Levinsen G. M. R. (1913) Systematic studies on the Sertulariidae. Videnskabelige Meddelelser fra Dansk Naturhistorik Forening i Kjøbenhavn 64, 249–323.

Marques A. C. (1996) A critical analysis of a cladistic study of the genus Eudendrium (Cnidaria: Hydrozoa), with some comments on the family Eudendriidae. Journal of Comparative Biology 1(3/4), 153–162.

Marques A. C. (2001) Simplifying hydrozoan classification: inappropriateness of the group Hydroidomedusae in a phylogenetic context. Contributions to Zoology 70(3), 175–179.

Marques A. C., Migotto A. E. (2001) Cladistic analysis and new classification of the family Tubulariidae (Hydrozoa, Anthomedusae). Papéis Avulsos de Zoologia 41(25), 465–488.

Marques A. C., Altuna A., Peña Cantero A. L., Migotto A. E. (2004) Redescription of Bedotella armata from the Biscay Bay, northern Spain, the type species of Bedotella Stechow, 1913 (Cnidaria, Hydrozoa, Lafoeidae), with comments on its taxonomic position. Hydrobiologia 530/531, 223–230.
CrossRef |

Migotto A. E. (1996) Benthic shallow-water hydroids (Cnidaria, Hydrozoa) of the coast of São Sebastião, Brazil, including a checklist of Brazilian hydroids. Zoologische Verhandelingen 306, 1–125.

Migotto A. E. (1998) The life cycle of Sertularia marginata Kirchenpauer, 1864 (Cnidaria, Hydrozoa): a medusoid-producing sertulariid. Journal of Natural History 32, 1–12.

Migotto A. E., Andrade L. P. (2000) The life cycle of Hebella furax (Cnidaria:Hydrozoa): a link between a lafoeid hydroid and a laodiceidae medusae. Journal of Natural History 34, 1871–1888.
CrossRef |

Migotto A. E., Marques A. C. (1999) Redescription of Dentitheca bidentata (Cnidaria: hydrozoa, Plumulariidae), with notes on its life cycle. Journal of Natural History 33, 949–960.
CrossRef |

Millard N. A. H. (1975) Monograph on the Hydroida of southern Africa. Annals of South African Museum 68, 1–513.

Millard N. A. H. (1977a) Hydroids from the Kerguelen and Crozet shelves, collected by the cruise MD.03 of the Marion-Dufresne. Annals of South African Museum 73(1), 1–47.

Millard N. A. H. (1977b) Hydroida. The South African Museum’s Meiring Naude cruises. Part 3. Annals of South African Museum 73(5), 105–131.

Naumov D. V. (1969). ‘Hydroids and Hydromedusae of the USSR.’ (Israel Program for Scientific Translation: Jerusalem, Israel.)

Naumov D. V., Stepan’yants S. D. (1962) Hydroida (Thecaphora) collected in the Antarctic and Subantarctic by the Soviet Antarctic Expedition with the M.V. “Ob,” in Antarctic and Subantarctic waters. Biological results of the Soviet Antarctic Expedition, 1955–1958 1, 68–106.

Nixon K. C., Carpenter J. M. (1993) On outgroups. Cladistics 9, 413–426.
CrossRef |

Nutting C. C. (1901a) The hydroids of the Woods Hole region. Bulletin of the United States Fish Commission 19, 325–386.

Nutting C. C. (1901b) Papers from the Harriman Alaska Expedition. XXI. The hydroids. Proceedings of the Washington Academy of Sciences 3, 157–216.

Nutting C. C. (1905) Hydroids of the Hawaiian Islands collected by the steamer Albatross in 1902. Bulletin of the United States Fish Commission 23(3), 931–959.

Nutting C. C. (1927) Report on Hydroida collected by the United States Fisheries steamer Albatross in the Philippine region, 1907–1910. Bulletin of the United States National Museum 100, 195–242.

Peña (Cantero) A. L., García-Carrascosa A. M. (1993) The coppinia of Abietinella operculata (Lafoeidae: Hydrozoa Leptomedusae) and its systematic position. Journal of Natural History 27(5), 1003–1011.

Peña Cantero A. L., García Carrascosa A. M. (2002) The benthic hydroid fauna of the Chafarinas Islands (Alborán Sea, western Mediterranean). Zoologische Verhandelingen 337, 1–180.

Peña Cantero A. L., Marques A. C. (1999) Phylogenetic analysis of the Antarctic genus Oswaldella Stechow, 1919 (Hydrozoa, Leptomedusae, Kirchenpaueriidae). Contributions to Zoology (Amsterdam, Netherlands : 1995) 68, 83–93.

Peña Cantero A. L., García Carrascosa A. M., Vervoort W. (1998) On the species of Filellum Hincks, 1868 (Cniaria: Hydrozoa) with the description of a new species. Journal of Natural History 32, 297–315.

Peña Cantero A. L., Vervoort W., Watson J. E. (2003) On Clathrozoellidae (Cnidaria, Hydrozoa, Anthoathecatae), a new family of rare deep-water leptolids, with the description of three new species. Zoologische Verhandelingen 345, 281–296.

Peña Cantero A. L., Marques A. C., Migotto A. E. (2004) Redescription of Acryptolaria normani Nutting, 1927 (Cnidaria, Hydrozoa, Lafoeidae), junior synonym of Zygophylax tizardensis Kirkpatrick, 1890 (Cnidaria, Hydrozoa, Lafoeidae). The Raffles Bulletin of Zoology 52(1), 1–6.

Quelch J. J. (1885) On some deep-sea and shallow water Hydrozoa. Annals and Magazine of Natural History 16(5), 1–20.

Ramil F., Vervoort W. (1992) Report on the Hydroida collected by the “BALGIM” expedition in and around the Strait of Gibraltar. Zoologische Verhandelingen 277, 3–262.

Rees W. J., Vervoort W. (1987) Hydroids from the John Murray Expedition to the Indian Ocean, with revisory notes on Hydrodendron, Abietinella, Cryptolaria and Zygophylax (Cnidaria: Hydrozoa). Zoologische Verhandelingen 237, 1–209.

Ritchie J. (1907) On collections of the Cape Verde Islands marine fauna, made by Cyril Crossland of St Andrews University, July to September 1904. The Hydroids. Proceedings of the Zoological Socitey of London 1907, 488–514.

Schuchert P. (2001) Hydroids of Greenland and Iceland (Cnidaria, Hydrozoa). Meddelselser om Grønland 53, 1–184.

Schuchert P. (2003) Hydroids (Cnidaria, Hydrozoa) of the Danish expedition to the Kei Islands. Steenstrupia 27(2), 137–256.

Spencer W. B. (1891) A new family of Hydroidea, together with a description of the structure of a new species of Plumularia. Transactions of the Royal Society of Victoria 2(1), 121–140.

Stechow E. (1909) Hydroidpolypen der japanischen Ostküste. 1. Theil. Athecata und Plumularidae. Abhandlungen der Mathematisch-Physikalischen Klasse der Königlichen Bayerischen Akademie der Wissenschaften (Supplement) 1(6), 1–111.

Stechow E. (1913) Neue Genera thecater Hydroiden aus der Familie der Lafoeiden und neue Species von Thecaten aus Japan. Zoologischer Anzeiger 43(3), 137–144.

Stechow E. (1921a) Neue Genera und Species von Hydrozoen und anderen Evertebraten. Archiv für Naturgeschichte [A]. 87(3), 248–265.

Stechow E. (1921b) Ueber Hydroiden der Deutschen Tiefsee-Expedition, nebst Bemerkungen über einige andre Formen. Zoologischer Anzeiger 53(9–10), 223–236.

Stechow E. (1922) Zur Systematik der Hydrozoen, Stromatoporen, Siphonophoren, Anthozoen und Ctenophoren. Archiv für Naturgeschichte [A]. 88(3), 141–155.

Stechow E. (1923) Zur Kenntnis der Hydroidenfauna des Mittelmeeres, Amerikas und anderer Gebiete. II. Teil. Zoologischer Jahrbücher. Abteilung für Systematik, Ökologie un Geographie der Tiere 47(1), 29–270.

Stechow E. (1925) Hydroiden der Deutschen Tiefsee-Expedition. Wissenschaftliche Ergebnisse de Deutschen Tiefsee-Expedition auf dem Dampfer “ Valdivia 1898–1899(17), 383–546.

Stepan’yants S. D. (1979) Gidroidy vod Antarktikh I Subantarktikh. Rezultaty Biologieskikh Issledovaniy Sovietskikh Antarkticheskikh Ekspedtsiy, Akademiya Nauk SSSR, Leningrad 22(30), 1–200.

Swofford D. L. (2001). ‘PAUP* Phylogenetic Analysis Using Parsimony version 4.0 b10.’ (Sinauer Associates: Sunderland, MA, USA.)

Totton A. K. (1930) Coelenterata. Part V. Hydroida. British Antarctic (“Terra Nova”) Expedition, 1910. Natural History Report, Zoology 5(5), 131–252.

Vanhöffen E. (1910) Die Hydroiden der Deutschen Südpolar-Expedition 1901–1903. Deutsche Südpolar-Expedition 1901–1903, Zoologie (4), 269–340.

Vervoort W. (1966) Bathyal and abyssal hydroids. Galathea Report 8, 97–173.

Vervoort W. (1972) Hydroids from the Theta, Vema and Yelcho cruises of the Lamont-Doherty geological observatory. Zoologische Verhandelingen 120, 1–247.

Vervoort W. (1987). Evaluation of taxonomic characters in the Hydroida, particularly in the Thecata (= Leptomedusae). In ‘Modern trends in the Systematics, Ecology and Evolution of Hydroids and Hydromedusae’. (Eds J. Bouillon, F. Boero, F. Cicogna and P. F. S. Cornelius.) pp. 83–103. (Clarendon Press: Oxford, UK.)

Vervoort W., Vasseur P. (1977) Hydroids from French Polynesia with notes on distribution and ecology. Zoologische Verhandelingen 159, 3–98.

Vervoort W., Watson J. E. (1996) A rare hydroid, Clathrozoella drygalskii (Vanhöffen, 1910) in the waters of Australia and New Zealand. Scientia Marina 60(1), 117–120.

Vervoort W., Watson J. E. (2003) The marine fauna of New Zealand: Leptothecata (Cnidaria: Hydrozoa) (Thecate Hydroids). NIWA Biodiversity Memoir 119, 1–538.

Watson J. E. (1969) Scoresbia, a new hydroid genus from South Australian waters. Transactions of the Royal Society of Southern Australia 93, 111–116.

Werner B. (1965) Die Nesselkapseln der Cnidaria, mit besonderer Berücksichtigung der Hydroida. 1. Klassification und Bedeutung für die Systematik und Evolution. Helgoländer Wissenschaftliche Meeresuntersuchungen 12, 1–39.
CrossRef |

Rent Article (via Deepdyve) Export Citation Cited By (6)