Invertebrate Systematics Invertebrate Systematics Society
Systematics, phylogeny and biogeography

Morphological phylogeny of the family Actinostolidae (Anthozoa : Actiniaria) with description of a new genus and species of hydrothermal vent sea anemone redefining the family Actinoscyphiidae

Estefanía Rodríguez A B , Christopher N. Castorani A and Marymegan Daly A
+ Author Affiliations
- Author Affiliations

A Department of Evolution, Ecology and Organismal Biology, Ohio State University, Columbus, OH 43212, USA.

B Corresponding author. Email:

Invertebrate Systematics 22(4) 439-452
Submitted: 23 October 2007  Accepted: 29 April 2008   Published: 14 October 2008


We describe a new genus and species of hydrothermal vent sea anemone from the eastern north Pacific Ocean. The combination of characters in Alvinactis reu, gen. et sp. nov. is unique among currently known genera of the actiniarian superfamily Mesomyaria; most notable among its external features is a belt of verrucae and cinclides in the distal column. We assess the placement of Alvinactis, gen. nov. and evaluate taxonomic features used to distinguish groups within Actinostolidae Carlgren, 1893 and Actinoscyphiidae Stephenson, 1920 with a cladistic analysis of morphological characters. Phylogenetic analysis reveals that Alvinactis, gen. nov. and several genera previously ascribed to Actinostolidae belong in Actinoscyphiidae. Morphological evidence fails to support monophyly of Actinostolidae, but does support monophyly of the previously proposed subfamily Actinostolinae.

Additional keywords: Actinoscyphiidae, Cnidaria, deep-sea diversity, hydrothermal vents, Pacific Ocean.


Specimens were collected by Janet Voight, with the assistance of the scientific party of cruise AT 11–03, the crew of the R/V Atlantis and the crew of the DSV Atlantis. J. Gerber of the FMNH helped in the accession and loan of the specimens. Collection of specimens was funded through NSF DEB-0072695 to J. Voight; their description was funded through NSF EF-0531763 to MD and a OSU CBS Dean’s Undergraduate Research award to CNC. J. Voight of the FMNH, S. Cairns of the USNM, R. Van Syoc of the CAS, and O. Tendal of the ZMUC provided comparative material. J. Wenzel gave advice on coding characters and identifying networks within the tree. A. Reft is thanked for her helpful advice and comments on cnidae.


Andres A. (1883). ‘Le Attinie (Monografía)’. (Coi Tipi der Salviucci: Roma.)

Bremer K. (1994) Branch support and tree stability. Cladistics 10, 295–304.
CrossRef |
open url image1

Carlgren O. (1893) Studien über Nordische Actinien. Kungliga Svenska Vetenskaps – Akademiens Handlingar 25, 1–148. open url image1

Carlgren O. (1899) Zoantharien. Hamburger Magalhaensische Sammelreise 4(1), 1–48. open url image1

Carlgren O. (1918) Die Mesenterienanordnung der Halcuriiden. Kungliga Fysiografiska Sällskapets Handlingar 29(29), 1–37. open url image1

Carlgren O. (1921) Actiniaria. I. Danish Ingolf – Expedition 5(9), 1–241. open url image1

Carlgren O. (1924) Actiniaria from New Zealand and its Subantarctic Islands. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening (Copenhagen) 77, 179–261. open url image1

Carlgren O. (1925) Zur Kenntnis der Hexacorallen. Zoologischer Anzeiger 65, 87–99. open url image1

Carlgren O. (1927) Actiniaria and Zoantharia. Further Zoological Results of the Swedish Antarctic Expedition 1901–1903 2(3), 1–102. open url image1

Carlgren O. (1928) Actiniaria der Deutschen Tiefsee-Expedition. Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expedition auf dem Dampfer “Valdivia” 1898–1899 4(22), 125–266. open url image1

Carlgren O. (1932). Die Ceriantharien, Zoantharien und Actiniarien des arktischen Gebietes. In ‘Eine Zusammenstellung der arktischen Tierformen mit besonderer Berücksichtigung des Spitzbergen-Gebietes auf Grund der Ergebnisse der Deutschen Expedition in das Nördliche Eismeer im Jahre 1898. Vol. 6’. (Eds F. Römer, F. Schaudinn, A. Brauer and W. Arndt.) pp. 255–266. (Gustav Fischer: Jena.)

Carlgren O. (1949) A survey of the Ptychodactiaria, Corallimorpharia and Actiniaria. Kungliga Svenska Vetenskaps – Akademiens Handlingar 1, 1–121. open url image1

Carlgren O. (1956) Actiniaria from depths exceeding 6000 meters. Galathea Reports 2, 9–16. open url image1

Daly M. (2004) Anatomy and taxonomy of three species of sea anemones (Cnidaria: Anthozoa: Actiniidae) from the Gulf of California, including Isoaulactina hespervolita Daly, n. sp. Pacific Science 58, 377–390.
CrossRef |
open url image1

Daly M., Gusmão L. (2007) The first sea anemone (Cnidaria, Anthozoa) from a whale fall. Journal of Natural History 41, 1–11.
CrossRef |
open url image1

Daly M., Chaudhuri A., Gusmão L., Rodríguez E. (2008) Phylogenetic Relationships among sea anemones (Cnidaria: Anthozoa: Actiniaria). Molecular Phylogenetics and Evolution 48, 292–301.
CrossRef | PubMed | open url image1

Danielssen D. C. (1890). Actinida. In ‘Den Norske Nordhavs-Expedition 1876–1878. Zoology’. p. 184. (Grøndahl and Søn: Christiania.)

den Hartog J. C. (1987) A redescription of the sea anemone Bunodosoma biscayensis (Fischer, 1874) (Actiniaria, Actiniidae). Zoologische Mededeelingen 61, 533–559. open url image1

Doumenc D., Van-Präet M. (1988) Actinies abyssales d’un site hydrothermal du Pacifique oriental. Oceanologica Acta 8, 61–68. open url image1

Edmands S., Fautin D. G. (1991) Redescription of Aulactinia veratra n. comb. (=Cnidopus veratra) (Coelenterata: Actiniaria) from Australia. Records of the Western Australian Museum (Perth) 15(1), 59–68. open url image1

England K. W. (1991) Nematocysts of sea anemones (Actiniaria, Ceriantharia, and Corallimorpharia: Cnidaria): nomenclature. Hydrobiologia 216/217, 691–697.
CrossRef |
open url image1

Fautin D. G. (1988). Importance of nematocysts to Actinian taxonomy. In ‘The Biology of Nematocysts’. (Eds D. A. Hessinger and H. M. Lenhoff.) pp. 487–500. (Academic Press: San Diego.)

Fautin D. G. (2007). ‘Hexacorallians of the World.’ Available online at [Accessed 18 December 2007].

Fautin D. G., Barber B. R. (1999) Maractis rimicarivora, a new genus and species of sea anemone (Cnidaria: Anthozoa: Actiniaria: Actinostolidae) from an Atlantic hydrothermal vent. Proceedings of the Biological Society of Washington 112, 624–631. open url image1

Fautin D. G., Hessler R. R. (1989) Marianactis bythios, a new genus and species of actinostolid sea anemone (Coelenterata: Actiniaria) from the Mariana vents. Proceedings of the Biological Society of Washington 102, 815–825. open url image1

Goloboff P. A. (1999). NONA (software and documentation). Available at [Accessed 20 October 2007].

Gosse P. H. (1859) Characters and descriptions of some new British sea anemones. Annals and Magazine of Natural History 3(13), 46–50. open url image1

Hertwig R. (1882). ‘Die Actinien der Challenger expedition.’ (Gustav Fischer: Jena.)

López-González P. J. , and Segonzac M. (2006). Cnidaria: Anthozoa: Actiniaria: Actinostolidae. In ‘Handbook of Deep-Sea Hydrothermal Vent Fauna’. (Eds D. Desbruyeres, M. Segonzac, and M. Bright.) pp. 544. (Denisa: Linz, Austria.)

López-González P. J., Rodríguez E., Segonzac M. (2005) A new species of sea anemone (Cnidaria: Anthozoa: Actiniaria) from Manus Basin Hydrothermal vents, South-western Pacific. Marine Biology Research 1, 326–337.
CrossRef |
open url image1

López-González P. J., Rodríguez E., Gili J. M., Segonzac M. (2003) New records on sea anemones (Anthozoa: Actiniaria) from hydrothermal vents and cold seeps. Zoologische Verhandelingen Leiden 345, 215–243. open url image1

Mariscal R. N. (1974). Nematocysts. In ‘Coelenterate biology: reviews and new perspectives’. (Eds L. Muscatine and H. M. Lenhoff.) pp. 129–178. (Academic Press: New York.)

McMurrich J. P. (1893) Report on the Actiniæ collected by the United States Fish Commission Steamer Albatross during the winter of 1887–1888. Proceedings of the United States National Museum 16, 119–216. open url image1

Milne Edwards H. (1857). ‘Histoire Naturelle des Coralliaires ou Polypes Proprement Dits. Vol. 1.’ (Librairie Encyclopédique de Roret: Paris.)

Milne Edwards H., Haime J. (1851) Reserches sur les polypiers; septième mémoire. Monographie des poritides. Annales des Sciences Naturelles 16(3), 21–70. open url image1

Nixon K. C. (1999). ‘Winclada, version 0.9.9.’ Available at [Accessed 20 October 2007].

Östman C. (2000) A guideline to nematocyst nomenclature and classification, and some note on the systematic value of nematocysts. Scientia Marina 64(1), 31–46. open url image1

Presnell J. K. , and Schreibman M. P. (1997). ‘Humason’s animal tissue techniques.’ pp. 572. (The Johns Hopkins University Press: Baltimore.)

Riemann-Zürneck K. (1978a) Tiefsee-Aktinien der familie Actinoscyphiidae aus dem Nordatlantik (Actiniaria, Mesomyaria). Zoologica Scripta 7, 145–153.
CrossRef |
open url image1

Riemann-Zürneck K. (1978b) Actiniaria des Südwestatlantik IV. Actinostola crassicornis (Hertwig, 1882) mit einer Diskussion verwandter Arten. Veröffentlichungen des Institutes für Meeresforschung Bremerhaven 17, 65–85. open url image1

Riemann-Zürneck K. (1997) Anthosactis janmayeni Danielssen, 1890, a rare high-arctic sea anemone. Polar Biology 17, 487–491.
CrossRef |
open url image1

Sanamyan N. P., Sanamyan K. E. (2007) Deep-water Actiniaria from East Pacific hydrothermal vents and cold seeps. Invertebrate Zoology 4(1), 83–102. open url image1

Schmidt H. (1969) Die Nesselkapseln der Aktinien und ihre differentialdiagnostiche Bedeutung. Helgoländer Wissenschaftliche Meeresuntersuchungen 19, 284–317.
CrossRef |
open url image1

Schmidt H. (1972) Die Nesselkapseln der Anthozoen und ihre Bedeutung fur die phylogenetische Systematik. Helgoländer Wissenschaftliche Meeresuntersuchungen 23, 422–458.
CrossRef |
open url image1

Schmidt H. (1974). On evolution in the Anthozoa. In ‘Proceedings of the Second International Coral Reef Symposium, Vol. 1’. (Eds A. M. Cameron et al.) pp. 533–560. (Great Barrier Reef Committee: Brisbane, Australia.)

Stephenson T. A. (1918) Coelenterata. I. Actiniaria. Natural History Reports on British Antarctic (“Terra Nova”) Expedition 1910 5(1), 1–68. open url image1

Stephenson T. A. (1920) On the classification of Actiniaria. Part I. Forms with acontia and forms with a mesogleal sphincter. The Quarterly Journal of Microscopical Science 64, 425–574. open url image1

Stephenson T. A. (1921) On the classification of Actiniaria. Part II. Consideration of the whole group and its relationships, with special reference to forms not treated in Part I. The Quarterly Journal of Microscopical Science 65, 493–576. open url image1

Stephenson T. A. (1928). ‘The British Sea Anemones. Vol. 1.’ (The Ray Society: London.)

Verrill A. E. (1869) Review of the corals and polyps of the west coast of America. Transactions of the Connecticut Academy of Arts and Sciences 1(6), 377–558. open url image1

Verrill A. E. (1883) Reports on the Anthozoa, and on some additional species dredged by the “Blake” in 1877–1879, and by the U. S. Fish Commission Steamer “Fish Hawk” in 1880–82. Bulletin of the Museum of Comparative Zoology [Harvard University] 11(1), 1–72. open url image1

Verrill A. E. (1899) Descriptions of imperfectly known and new Actinians, with critical notes on other species, IV. American Journal of Science and Arts 7(4), 205–218. open url image1

White T. R., Wakefield Pagels A. K., Fautin D. G. (1999) Abyssal sea anemones (Cnidaria: Actiniaria) of the northeast Pacific symbiotic with molluscs: Anthosactis nomados, a new species, and Monactis vestita (Gravier, 1918). Proceedings of the Biological Society of Washington 112(4), 637–651. open url image1

Williams R. B. (1998) Measurements of cnidae from sea anemones (Cnidaria: Actiniaria), II: further studies of differences amongst sample means and their taxonomic relevance. Scientia Marina 62, 361–372. open url image1

Williams R. B. (2000) Measurements of cnidae from sea anemones (Cnidaria: Actiniaria), III: ranges and other measures of statistical dispersion, their interrelations and taxonomic relevance. Scientia Marina 64, 49–68. open url image1

Appendix 1.  Morphological characters used in cladistic analysis of Actinostolidae
Characters in bold used in Carlgren’s 1949 key to the actinostolid genera. Characters that do not manifest exclusive states in all members of a taxon have been broken into several binary characters (e.g. #s 22 and 23) rather than coded as single multistate characters. Characters applicable only to outgroup taxa indicated
Click to zoom

Appendix 2.  Morphological character state distributions for outgroup and ingroup genera
Dash indicates that the corresponding state is unknown or inapplicable. Outgroup genera in bold. See Appendix 1 for character list
Click to zoom

Rent Article (via Deepdyve) Export Citation Cited By (9)