Skip to main content Skip to main navigation menu Skip to site footer
Type: Article
Published: 2026-04-02
Page range: 53-81
Abstract views: 36
PDF downloaded: 2

On the type specimens of Parazoanthus axinellae (Schmidt, 1862) (Hexacorallia: Zoanthidea) and its subspecies

Evolution of Marine Symbioses Laboratory; Department of Marine and Environmental Science; Nova Southeastern University; Fort Lauderdale; Florida; USA
Department of Biology; Faculty of Science University of Zagreb; Zagreb; Croatia
Musée Océanographique de Monaco; Monaco
Musée Océanographique de Monaco; Monaco
Department of CAPE; Stazione Zoologica Anton Dohrn di Napoli; Naples; Italy
Centre of Natural History; Universalmuseum Joanneum; Graz; Austria
UMR7044 Archimède du CNRS; Musée Zoologique de Strasbourg; Strasbourg; France
Reef Ecology; Ecosystems & Functions Laboratory; Department of Marine and Environmental Science; Nova Southeastern University; Fort Lauderdale; Florida; USA; National Coral Reef Institute; Halmos College of Arts and Sciences; Nova Southeastern University; Fort Lauderdale; Florida; USA
Evolution of Marine Symbioses Laboratory; Department of Marine and Environmental Science; Nova Southeastern University; Fort Lauderdale; Florida; USA; National Coral Reef Institute; Halmos College of Arts and Sciences; Nova Southeastern University; Fort Lauderdale; Florida; USA
Department of CAPE; Stazione Zoologica Anton Dohrn di Napoli; Naples; Italy
Evolution of Marine Symbioses Laboratory; Department of Marine and Environmental Science; Nova Southeastern University; Fort Lauderdale; Florida; USA; National Coral Reef Institute; Halmos College of Arts and Sciences; Nova Southeastern University; Fort Lauderdale; Florida; USA
Coelenterata anatomy natural history collections marginal muscle symbiosis taxonomy

Abstract

Subspecies of Parazoanthus axinellae have been largely ignored since their inception and modern authors write of morphotypes rather than attempting to assign their specimens to these existing taxa. This hesitation to apply subspecies names is rooted in taxon definitions written to 19th century standards coupled with failure to designate type specimens that could be reexamined for clarification. Revealing the types or verifying that they no longer exist so that topotypes may be designated as neotypes creates a path for clarification of existing taxonomic hypotheses and discovering which modern morphotypes or cryptic taxa lay outside those circumscriptions. Here we reconstructed the history and updated the diagnoses of the types of P. axinellae and its nominal subspecies by reviewing historic scientific and archival documents, verifying type existence with their current caretakers, and reexamining their microanatomy. We identified type specimens and designated lectotypes and paralectotypes from the syntype series of Parazoanthus axinellae and subspecies axinellae, liguricus, and muelleri, redescribed them from histology of their type material, and provide diagnostic traits for identification in the field or laboratory. Type material for brevitentacularis was not recovered and permission to resample the type location has been denied.

 

References

  1. Abel, E. (1955) Eine neue Krustenanemone der Adria, Epizoanthus paxi, nov. spec. Österreichische Zoologische Zeitschrift, 6, 1–6.
  2. Abel, E. (1959) Zur Kenntnis der marinen Höhlenfauna unter besonderer Berücksichtigung der Anthozoen. Pubblicazioni della Stazione Zoologica di Napoli, 30, 1–94.
  3. Appeltans, W., Ahyong, S.T., Anderson, G., Angel, M.V., Artois, T., Bailly, N., Bamber, R., Barber, A., Bartsch, I., Berta, A., Błażewicz-Paszkowycz, M., Bock, P., Boxshall, G., Boyko, C.B., Brandão, S.N., Bray, R.A., Bruce, N.L., Cairns, S.D., Chan, T.-Y., Cheng, L., Cirino, R.N., Collins, A.G., Cornelius, P.F., Couwberg, N., Cowman, P.F., Cruz-da-Silva, F., Cummins, V., Dadahouh-Guebas, F., Davison, I., De Broyer, C., Dekeyzer, S., Desiderato, A., Di Capua, I., Dolmer, P., Dionisios, E., Eibye-Jacobsen, D., Enriquez, S., Fautin, D.G., Feist, S.W., Fransen, C.H., Furuya, H., Garcia-Alvarez, O., Gerken, S., Gibson, D.I., Gittenberger, A., Gofas, S., Gómez-Daglio, L., Williams, G.C., Göthel, H., Guiry, M.D., Hernandez, F., Hoeg, J.T., Hillewaert, H., Hojsgaard, P., Huys, R., Ishimaru, S., Jamsari, A.J., Janson, G.S., Janssen, R., Javidpour, J., Jiang, J.-C., Jozwiak, P., Kamiya, T., Kasyan, V.V., Kawai, H., Kim, J., King, G.S., Kirk, P.M., Knight-Jones, P., Koenemann, S., Kolb, J., Kotov, A.A., Kroh, A., Lützen, J., Mackelfresh, S.J., Madin, L.P., Mah, C.L., Mallik, J.G., Malyutina, M.V., Mann, S.J., Marshall, B.A., Matsuura, K., McInnes, S.J., Mees, J., Meland, K., Miller, C.G., Mills, C.E., Molodtsova, T.N., Mooi, R., Morandini, A.C., Naruse, T., Nielsen, C., Norenburg, J.L., Opresko, D.M., Osawa, M., Paulay, G., Paxton, H., Podchiplov, V.G., Polaszek, A., Polz, M.F., Reimer, J.D., Rius, M., Rocha, R.M., Saiz-Salinas, J.I., Scarabino, V., Schander, C., Schloss, S.J., Schotte, M., Segers, H., Sørensen, M.V., Sluys, R., Smith, A.B., Swalla, B.J., Tchesunov, A.V., Taiti, S., Tokuda, G., Tomikawa, K., van der Land, J., van Soest, R.W., van Tussenbroek, B.I., Vanaverbeke, J., Walker-Smith, G.K., Walter, T.C., Warren, A., Williams, G.C., Wilson, S.P., van Winston, J.E., Wyatt, A.S., Yamasaki, H., Yoo, J.-S., Zenetos, A., Zhan, Z., Zhadan, A.E., Zelazny, B. & Costello, M.J. (2012) The Magnitude of Global Marine Species Diversity. Current Biology, 22, 2189–2202. https://doi.org/10.1016/j.cub.2012.09.036
  4. Blainville, H.d. (1830) Zoophytes. In: Cuvier, G.-F. (Ed.), Dictionnaire des Sciences Naturelles, dans lequel on traite méthodiquement des différens êtres de la nature, considérés soit en eux-mêmes, d’après l’état actuel de nos connoissances, soit relativement a l’utilité qu’en peuvent retirer la médecine, l’agriculture, le commerce et les arts. Suivi d’une biographie des plus célèbres naturalistes. Ouvrage destiné aux médecins, aux agriculteurs, aux commercans, aux artistes, aux manufacturiers, et à tous ceux qui ont intérêt à connoître les productions de la nature, leurs caractères génériques et spécifiques, leur lieu natal, leurs propriétés et leurs usages. F. G. Levrault, Strasbourg et Paris/Le Normant, Paris, pp. 1–546.
  5. Bourne, G.C. (1900) The Anthozoa. In: Lankester, E.R. (Ed.), A Treatise on Zoology. Part II. The Porifera and Coelentera. Adam & Charles Black, London, pp. 1–84.
  6. Bowerbank, J.S. (1858) On the anatomy and physiology of the Spongiadae. Part I. On the Spicula. Philosophical transactions of the Royal Society of London, 148, 279–332. https://doi.org/10.1098/rstl.1858.0016
  7. Cachet, N., Genta-Jouve, G., Ivanisevic, J., Chevaldonné, P., Sinniger, F., Thomas, O.P. & Culioli, G. (2015) Metabolomic profiling reveals deep chemical divergence between two morphotypes of the zoanthid Parazoanthus axinellae. Scientific Reports, 5, 8282. https://doi.org/10.1038/srep08282
  8. Carlgren, O. (1935) Di alcune Attinie e Zoantari raccolti nel Golfo di Genova. Bollettino dei musei e laboratorii di zoologia e anatomia comparata della R. Università di Genova, 15, 3–14.
  9. Cerrano, C., Totti, C., Sponga, F. & Bavestrello, G. (2006) Summer disease in Parazoanthus axinellae (Schmidt, 1862)(Cnidaria, Zoanthidea). Italian Journal of Zoology, 73, 355–361. https://doi.org/10.1080/11250000600911675
  10. Cinar, M.E., Yokeş, M.B., Cinar, S.A. & Bakir, A.K. (2014) Checklist of Cnidaria and Ctenophora from the coasts of Turkey. Turkish Journal of Zoology, 38, 677–697. https://doi.org/10.3906/zoo-1405-68
  11. Colbert, J., Hellenbrand, H., Swain, A.K. & Swain, T.D. (2026) Revision of Hawaiian seamount Zoanthidea genera to bridge the parataxonomy gap, retain Gerardiidae, and introduce Antipathozoanthidae nov. fam. and Vitrumanthidae nov. fam. Marine Biovidersity. [in press]
  12. Desqueyroux-Faúndez, R. & Stone, S. (1992) O. Schmidt sponge catalogue: an illustrated guide to the Graz Museum Collection, with notes on additional material. Muséum d’histoire naturelle, Geneva, 190 pp., XLIX pls.
  13. Enrichetti, F., Dominguez-Carrió, C., Toma, M., Bavestrello, G., Betti, F., Canese, S. & Bo, M. (2019) Megabenthic communities of the Ligurian deep continental shelf and shelf break (NW Mediterranean Sea). PLoS One, 14, e0223949. https://doi.org/10.1371/journal.pone.0223949
  14. Esper, E.J.C. (1794) Die Pflanzenthiere in Abbildungen nach der Natur mit Farben erleuchtet, nebst Beschreibungen. Vol. 1. Raspischen Buchhandlung, Nürnberg, xii + 320 pp.
  15. Evenhuis, N.L. (2008) A compendium of zoological type nomenclature: a reference source. No. 41. Bishop Museum Press, Honolulu, Hawaii, 23 pp.
  16. Gomes-Pereira, J.N., Vasco-Rodrigues, N., Afonso, A., Albuquerque, M., Berecibar, E., Bernardes, L., Calado, A., Castanheira, A., Tojeira, I., Tempera, F., Dias, F.C., Santos, R.S. & Campos, A.S. (2015) Mono-specific facies of Parazoanthus axinellae in Luiz Saldanha Marine Park. Marine Biodiversity Records, 8, e99. https://doi.org/10.1017/S1755267215000597
  17. Gómez-Gras, D., Linares, C., de Caralt, S., Cebrian, E., Frleta-Valić, M., Montero-Serra, I., Pagès-Escolà, M., López-Sendino, P. & Garrabou, J. (2019) Response diversity in Mediterranean coralligenous assemblages facing climate change: Insights from a multispecific thermotolerance experiment. Ecology and Evolution, 9, 4168–4180. https://doi.org/10.1002/ece3.5045
  18. Gray, J. (1832) Zoantharia. In: Woodfall, G. (Ed.), Synopsis of the contents of the British Museum. 26th Edition. G. (George) Woodfall, London, pp. 94–106.
  19. Gray, J. (1870) Catalogue of the sea-pens or Pennatulariidae in the collection of the British Museum. Trustees of the British Museum, London, 40 pp. https://doi.org/10.5962/bhl.title.11307
  20. Haddon, A.C. & Shackleton, A.M. (1891) A Revision of the British Actiniæ. Part II: The Zoantheæ. Scientific Transactions of the Royal Dublin Society, 4, 609–672. https://doi.org/10.5962/bhl.title.46280
  21. Haeckel, E. (1896) Systematische Phylogenie der Wirbellosen Thiere (Invertebrata). Zweiter Theil. Des Entwurfs einer systematischen Stammesgeschichte. Georg Reimer, Berlin, xviii + 720 pp. https://doi.org/10.1515/9783111443935
  22. Herberts, C. (1972) Étude systématique de quelques zoanthaires tempérés et tropicaux. Tethys, Supplemental 3, 69–156.
  23. Hertwig, R. (1882) Die Actinien der Challenger Expedition. Gustav Fischer, Jena, 119 pp.
  24. Hoppe, B. (2001) Pax, Ferdinand. In: Neue Deutsche Biographie. Vol. 20. Duncker & Humblot, Berlin, pp. 143–145.
  25. ICZN (1999) International Code of Zoological Nomenclature. 4th Edition. The International Trust for Zoological Nomenclature, London, xxix + 306 pp.
  26. Kise, H., Santos, M.E.A., Fourreau, C.J.L., Iguchi, A., Goto, R. & Reimer, J.D. (2023) Evolutionary patterns of host switching, lifestyle mode, and the diversification history in symbiotic zoantharians. Molecular Phylogenetics and Evolution, 182, 107732. https://doi.org/10.1016/j.ympev.2023.107732
  27. Manuel, R.L. (1988) British Anthozoa (Coelenterata: Octocorallia & Hexacorallia). In: Kermack, D.M. & Barnes, R.S.K. (Eds.), Synopses of British Fauna. The Linnean Society of London and The Estuarine and Brackish-water Sciences Association, Leiden, pp. 1–83. https://doi.org/10.1163/9789004627543
  28. Markovich, A. (2021) Applying integrative systematics to long-standing species boundary questions in the Zoanthidea. Master’s Thesis, Nova Southeastern University, ort Lauderdale-Davie, Florida. Available from: https://nsuworks.nova.edu/hcas_etd_all/70 (accessed 9 February 2026)
  29. McFadden, C.S., Quattrini, A.M., Brugler, M.R., Cowman, P.F., Dueñas, L.F., Kitahara, M.V., Rodriguez, E., Reimer, J.D. & Daly, M. (2021) Phylogenomics, Origin and Diversification of Anthozoans (Phylum Cnidaria). Systematic Biology, 70 (4), 635–647. https://doi.org/10.1093/sysbio/syaa103
  30. Montagu, G. (1814) An essay on sponges, with descriptions of all the species that have been discovered on the coast of Great Britain. Memoirs of the Wernerian Natural History Society, 2, 67–122.
  31. Montenegro, J., Fromont, J., Richards, Z., Kise, H., Gomez, O., Hoeksema, B.W. & Reimer, J.D. (2024) Museum collections as untapped sources of undescribed diversity of sponge-zoantharian associations with the description of six new species of Umimayanthus (Zoantharia: Parazoanthidae) from Western Australia and eastern Indonesia. Contributions to Zoology, 1, 1–57. https://doi.org/10.1163/18759866-bja10069
  32. Nardo, G.D. (1833) Auszug aus einem neuen System der Spongiarien, wonach bereits die Aufstellung in der Universitäts-Sammlung zu Padua gemacht ist. In: Oken, J. (Ed.), Isis, oder Encyclopädische Zeitung Coll. Oken, Jena, pp. 519–523.
  33. Ocaña, O., Moro, L., Herrera, R., Çinar, M.E., Brito, A., Cuervo, J.S. & Herrero, R. (2019) Parazoanthus axinellae: a species complex showing different ecological requierements. Revista de la Academia Canaria de Ciencias: Folia Canariensis Academiae Scientiarum, 31, 15–36.
  34. Pallas, P.S. (1766) Elenchus zoophytorum sistens generum adumbrationes generaliores et specierum cognitarum succintas descriptiones, cum selectis auctorum synonymis. Fransiscum Varrentrapp, Hague, 451 pp. https://doi.org/10.5962/bhl.title.6595
  35. Pax, F. (1937) Die Korallenfauna der Adria. Teil 1: Krustenanemonen. Thalassia, 2, 3–66.
  36. Pax, F. (1952) Die Antipatharien, Zoantharien und Actiniarien der “Hvar”-Expedition. Izvjesca-Reports, 6, 3–24.
  37. Pax, F. (1957) Die Zoantharien des Golfes von Neapel. Pubblicazioni della Stazione Zoologica di Napoli, 30, 309–329.
  38. Pax, F. & Lochter, H. (1935) Epizoanthus vatovai, eine neue Carcinoecien bildende Krustenanemone der Adria. Notizen des Deutsch-Italienischen Instituts für Meeresbiologie in Rovigno, 1 (18), 3–16.
  39. Pax, F. & Müller, I. (1955) Gli Antozoi del Museo Civico di Storia Naturale di Trieste Parte I: Antipatharia, Ceriantharia, Zoantharia, Actiniaria, Alcyonaria e Pennatularia. Atti del Museo Civico di Storia Naturale Trieste, 20, 103–129.
  40. Pax, F. & Müller, I. (1956a) La collection de Zoanthaires du Musée océanographique de Monaco. Bulletin de l’Institut océanographique, 53, 1–27.
  41. Pax, F. & Müller, I. (1956b) Westafrikanische Zoantharien. Bulletin de l’Institut Royal des Sciences Naturelles de Belgique, 32, 1–11.
  42. Pax, F. & Müller, I. (1957) Zoantharien des zoologischen museums in Genua. Ann del Mus Civ di Stor Nat Giacomo Doria, 69, 19–30.
  43. Pax, F. & Müller, I. (1962) Die anthozoenfauna der Adria. Fauna et Flora Adriatica. Vol. 3. Institut za Oceanografiju i Ribastvo Split, Split, 343 pp.
  44. Riedl, R. (1966) Biologie der Meereshöhlen: Topographie, Faunistik und Ökologie eines unterseeischen Lebensraumes: Eine Monographie. Verlag Paul Parey, Hamburg, 636 pp.
  45. Rosales Ruiz, A., Ocaña, O., de la Herrán, R., Navajas-Pérez, R., Ruiz Rejón, C., Congil Ross, A. & Ruiz Rejón, M. (2025) Going deep into Parazoanthus axinellae (Anthozoa: Zoantharia) complex: description of two species in the Alboran Sea based on an integrative approach. Marine Biodiversity, 55, 15. https://doi.org/10.1007/s12526-024-01493-x
  46. Schmidt, O. (1862) Die Spongien des adriatischen Meeres. Verlag Von Wilhelm Engelmann, Leipzig, viii + 88 pp., 7 pls.
  47. Swain, T.D. (2009) Phylogeny-based species delimitations and the evolution of host associations in symbiotic zoanthids (Anthozoa, Zoanthidea) of the wider Caribbean region. Zoological Journal of the Linnean Society, 156, 223–238. https://doi.org/10.1111/j.1096-3642.2008.00513.x
  48. Swain, T.D. (2010) Evolutionary transitions in symbioses: dramatic reductions in bathymetric and geographic ranges of Zoanthidea coincide with loss of symbioses with invertebrates. Molecular Ecology, 19, 2587–2598. https://doi.org/10.1111/j.1365-294X.2010.04672.x
  49. Swain, T.D. (2018) Revisiting the phylogeny of Zoanthidea (Cnidaria: Anthozoa): Staggered alignment of hypervariable sequences improves species tree inference. Molecular Phylogenetics and Evolution, 118, 1–12. https://doi.org/10.1016/j.ympev.2017.09.008
  50. Swain, T.D., Schellinger, J.L., Strimaitis, A.M. & Reuter, K.E. (2015) Evolution of anthozoan polyp retraction mechanisms: convergent functional morphology and evolutionary allometry of the marginal musculature in order Zoanthidea (Cnidaria: Anthozoa: Hexacorallia). BMC Evolutionary Biology, 15, 123. https://doi.org/10.1186/s12862-015-0406-1
  51. Swain, T.D., Strimaitis, A.M., Reuter, K.E. & Boudreau, W. (2017) Towards integrative systematics of Anthozoa (Cnidaria): evolution of form in the order Zoanthidea. Zoologica Scripta, 46, 227–244. https://doi.org/10.1111/zsc.12195
  52. Swain, T.D. & Swain, L.M. (2014) Molecular parataxonomy as taxon description: examples from recently named Zoanthidea (Cnidaria: Anthozoa) with revision based on serial histology of microanatomy. Zootaxa, 3796 (1), 81–107. https://doi.org/10.11646/zootaxa.3796.1.4
  53. Swain, T.D. & Wulff, J.L. (2007) Diversity and specificity of Caribbean sponge-zoanthid symbioses: a foundation for understanding the adaptive significance of symbioses and generating hypotheses about higher-order systematics. Biological Journal of the Linnean Society, 92, 695–711. https://doi.org/10.1111/j.1095-8312.2007.00861.x
  54. Terzin, M., Villamor, A., Marincich, L., Matterson, K., Paletta, M.G., Bertuccio, V., Bavestrello, G., Benedetti Cecchi, L., Boscari, E., Cerrano, C., Chimienti, G., Congiu, L., Fraschetti, S., Mastrototaro, F., Ponti, M., Sandulli, R., Turicchia, E., Zane, L., Abbiati, M. & Costantini, F. (2024) 2bRAD reveals fine-scale genetic structuring among populations within the Mediterranean zoanthid Parazoanthus axinellae (Schmidt, 1862). Coral Reefs, 43, 357–370. https://doi.org/10.1007/s00338-023-02456-7
  55. Topsent, E. (1934) Éponges observées dans les parages de Monaco. 1er Partie. Bulletin de l’Institut océanographique de Monaco, 650, 1–42.
  56. Vafidis, D. & Koukouras, A. (1998) Antipatharia, Ceriantharia and Zoantharia (Hexacorallia, Anthozoa) of the Aegean Sea with a checklist of the Mediterranean and Black Sea species. Annales de l’Institut Oceanographique, 74 (2), 115–126.
  57. Villamor, A., Signorini, L.F., Costantini, F., Terzin, M. & Abbiati, M. (2020) Evidence of genetic isolation between two Mediterranean morphotypes of Parazoanthus axinellae. Scientific Reports, 10, 13938. https://doi.org/10.1038/s41598-020-70770-z
  58. Zavodnik, D., Zavodnik, N. & Iveša, L. (2001) The 110th anniversary of the marine research station at Rovinj (Adriatic Sea, Croatia). Reference collections. Natura Croatica: Periodicum Musei Historiae Naturalis Croatici, 10, 53–60.

How to Cite

Markovich, A., Bakran-Petricioli, T., Baltzinger, E., Bruni, M., Cenci, E., Gunczy, J., Meister, M., Ploner, M., Swain, A.K., Travaglini, A. & Swain, T.D. (2026) On the type specimens of Parazoanthus axinellae (Schmidt, 1862) (Hexacorallia: Zoanthidea) and its subspecies. Zootaxa, 5787 (1), 53–81. https://doi.org/10.11646/zootaxa.5787.1.3