Dendrodorididae (Heterobranchia, Nudibranchia) from Persian Gulf with a description of a new species of Doriopsilla and remarks on the family

  • Fatemeh Maniei Leibniz Institute for the Analysis of Biodiversity Change, Museum Koenig Bonn, Adenauerallee 160, 53113 Bonn, Germany
  • Heike Wägele Leibniz Institute for the Analysis of Biodiversity Change, Museum Koenig Bonn, Adenauerallee 160, 53113 Bonn, Germany
Keywords: Dendrodoris, Iran, histology, species delimitation, haplotype networking


The family Dendrodorididae has a global distribution, with prevalence in tropical and subtropical intertidal zones. Three species of Dendrodorididae were collected from the intertidal zone of the northern coast of the Persian Gulf in Iran. Based on anatomical, histological, and molecular investigations they can be assigned to Dendrodoris fumata, Dendrodoris nigra, and a new species of Doriopsilla, D. aroni sp. nov. Molecular analyses of CO1 and 16S, including all genera of Dendrodorididae, members of the sister taxon Phyllidiidae, and other dorid outgroups resulted in a polyphyletic genus Dendrodoris, which is in contrast to the nuclear gene studies. Our molecular results confirm the differentiation between Dendrodoris rubra and D. fumataDendrodoris nigra, D. fumata, and D. krusensternii each consist of several clades, indicating cryptic species complexes requiring further investigation. We describe the presence of bacteria for the first time in the vestibular gland of D. fumata. Validation of the specimens of Doriopsilla from the Persian Gulf as a new species is supported by haplotype networking, genetic distance, and ABGD analyses of mitochondrial genes. Our CO1 analysis confirms a previous hypothesis that Cariopsilla is a junior synonym of Doriopsilla.


Abdollahi R., Darzi S.E., Rahimian H. & Naderloo R. 2020. Biodiversity of intertidal and shallow subtidal habitats in Abu Musa Island, Persian Gulf, Iran, with an updated species checklist. Regional Studies in Marine Science 33: 100975.

Alder J. & Hancock A. 1864. Notice of a collection of nudibranchiate Mollusca made in India by Walter Elliot Esq. with descriptions of several new genera and species. Transactions of the Zoological Society of London 5: 113–147.

Almada F., Levy A. & Robalo J. 2016. Not so sluggish: the success of the Felimare picta complex (Gastropoda, Nudibranchia) crossing Atlantic bio geographic barriers. PeerJ 4: e1561.

Amini-Yekta F. & Dekker H. 2021. An updated checklist of marine gastropods of the Persian Gulf and Gulf of Oman. Zootaxa 4957 (1): 1–71.

Amini-Yekta F., Asgari M. & Izadi S. 2016. Molluscs, Crustaceans and Echinoderms of the Intertidal Area of Qeshm Island (with Illustrated List of Other Invertebrates of this Region). Persian Gulf Biotechnology Park, Qeshm Island. [In Persian.]

Amini-Yekta F., Shokri M.R., Maghsoudlou A. & Rajabi-Maham H. 2019. Intertidal gastropod assemblages shaped by key environmental variables across the northern Persian Gulf and the Gulf of Oman. Marine Ecology 40 (3): e12545.

Astrin J.J. & Stüben P.E. 2008. Phylogeny in cryptic weevils: molecules, morphology and new genera of western Palaearctic Cryptorhynchinae (Coleoptera: Curculionidae). Invertebrate Systematics 22: 503–522.

Ávila C., Ballesteros M. & Ortea J. 1992. Una nueva especie de Doriopsilla Bergh, 1880 (Mollusca: Nudibranchia) del Archipiélago de Cabo Verde. Historia Animalium 1: 23–31.

Ballesteros M. 2023. “Resurrecting” species: the case of the sea slug “Dendrodoris temarana”. OPK Opistobranquis. Available from [accessed 17 Apr. 2023].

Bergh L.S.R. 1880. Die Doriopsen des Mittelmeeres. Jahrbücher der Deutschen Malakozoologischen Gesellschaft 7: 297–328. Available from [accessed 17 Apr. 2023].

Bergh L.S.R. 1902. I. Gasteropoda Opisthobranchiata. In: The Danish Expedition to Siam 1899–1900. Det Kongelige Danske Videnskabernes Selskabs Skrifter. 6 Raekke Naturvidenskabelig og Matematisk Afdeling 12 (2): 153–218. Available from [accessed 17 Apr. 2023].

Bergh L.S.R. 1907. The Opisthobranchiata of South Africa. Transactions of the South African Philosophical Society 17 (1): 1–144. [Dated 1908.] Available from [accessed 17 Apr. 2023].

Brodie G.D. 2001. Some comparative histological aspects of the dendrodorid genera Doriopsilla and Dendrodoris (Opisthobranchia: Nudibranchia). Bollettino Malacologico 37 (5–8): 99–104. Available from [accessed 17 Apr. 2023].

Brodie G.D., Willan R.C. & Collins J.D. 1997. Taxonomy and occurrence of Dendrodoris nigra and Dendrodoris fumata (Nudibranchia. Dendrodorididae) in the Indo-West Pacific Region. Journal of Molluscan Studies 63: 407–423.

Burn R. 1962. Notes on a collection of Nudibranchia (Gastropoda: Dorididae and Dendrodorididae) from South Australia with remarks on the species of Basedow and Hedley, 1905. Memoirs of the National Museum of Victoria 25: 149–172. https;//

Burn R. 1966. Notes on some opisthobranchs mainly from South Australia. Records of the South Australian Museum 15 (2): 329–352.

Clement M., Posada D. & Crandall K.A. 2000. TCS: a computer program to estimate gene genealogies. Molecular Ecology 9: 1657–1659.

Cunha T.J., Fernández-Simón J., Petrula M., Giribet G. & Moles J. 2023. Photographic checklist, DNA barcoding, and new species of sea slugs and snails from the Faafu Atoll, Maldives (Gastropoda: Heterobranchia and Vetigastropoda). Diversity 15: 219.

Edmunds M. 1971. Opisthobranchiate Mollusca from Tanzania (Suborder: Doridacea). Zoological Journal of the Linnean Society 50: 339–396.

Eisenbarth J.H., Undap N., Papu A., Schillo D., Dialao J., Reumschüssel S., Kaligis F., Bara R., Schäberle T.F., König G.M., Yonow N. & Wägele H. 2018. Marine Heterobranchia (Gastropoda, Mollusca) in Bunaken National Park, North Sulawesi, Indonesia – A follow-up diversity study. Diversity 10 (4): 127.

Eliot C.N.E. 1904. On some nudibranchs from East Africa and Zanzibar. Part III. Proceedings of the Zoological Society of London 1903 (2): 354–385. Available from [accessed 17 Apr. 2023].

Fatemi Y. & Attaran F.G. 2015. Checklist of the opisthobranchs (Heterobranchia: Gastropoda) along the Iranian coasts of the Gulf of Oman. Journal of Biodiversity and Environmental Sciences (JBES) 6 (3): 1–7.

Fatemi Y., Taheri M.R., Esmaeili H.R. & Shahdadi A. 2021. Morphological and molecular analysis of Dendrodoris fumata (Rüppell & Leuckart, 1830) (Gastropoda: Nudibranchia) from the Persian Gulf. Journal of Aquatic Ecology 10 (4): 49–57. Available from [accessed 17 Apr. 2023]. [In Persian.]

Furfaro G., Schreier Ch., Trainito E., Pontes M., Madrenas E., Girard P. & Mariottini P. 2022. The sea slug Doriopsilla areolata Bergh, 1880 (Mollusca, Gastropoda) in the Mediterranean Sea: another case of cryptic diversity. Diversity 14: 297.

Galià-Camps C., Cervera J.L., Valdés Á. & Ballesteros M. 2022. Attack on crypsis: molecular and morphological study of Dendrodoris Ehrenberg, 1831 (Mollusca: Gastropoda: Nudibranchia) from the Mediterranean Sea and Northern Atlantic Ocean reinstates Dendrodoris temarana Pruvot-Fol, 1953. Zootaxa 5133 (3): 383–406.

GBIF Secretariat 2023a. Dendrodoris krusensternii (Gray, 1850). GBIF Backbone Taxonomy. Available from [accessed 5 Oct. 2023].

GBIF Secretariat 2023b. Cariopsilla Ortea & Espinosa, 2006. GBIF Backbone Taxonomy. Available from [accessed 5 Sep. 2023].

Gohar H.A.F. & Soliman G.N. 1967. The biology and development of Dendrodoris (= Doridopsis) fumata (Rüppell and Leuckart) (Gastropoda, Nudibranchia). Publications of the Marine Biological Station Al-Ghardaqa, Red Sea 14: 31–54.

Goodheart J. & Valdés Á. 2013. Re-evaluation of the Doriopsilla areolata Bergh, 1880 (Mollusca: Opisthobranchia) subspecies complex in the eastern Atlantic Ocean and its relationship to South African Doriopsilla miniata (Alder & Hancock, 1864) based on molecular data. Marine Biodiversity 43: 113–120.

Gosliner T.M., Behrens D.W. & Valdés Á. 2008. Indo-Pacific Nudibranchs and Sea Slugs: A Field Guide to the World’s Most Diverse Fauna. Sea Challengers Inc., Monterey.

Gosliner T.M., Valdés Á. & Behrens D.W. 2018. Nudibranch and Sea Slug Identification: Indo-Pacific. Second edition. New World Publications, Jacksonville.

Hall T.A. 1999. BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series 41 (41): 95–98.

Hallas J.M., Chichvarkhin A. & Gosliner T.M. 2017. Aligning evidence: concerns regarding multiple sequence alignments in estimating the phylogeny of the Nudibranchia suborder Doridina. Royal Society. Open Science 5: 171095.

Hirose M., Hirose E. & Kiyomoto M. 2014. Identification of five species of Dendrodoris (Mollusca: Nudibranchia) from Japan, using DNA barcode and larval characters. Marine Biodiversity 45 (4): 769–780.

Huson D.H. & Scornavacca C. 2017. Dendroscope 3: an interactive tool for rooted phylogenetic trees and networks. Systematic Biology 61: 1061–1067.

Katoh K., Misawa K., Kuma K. & Miyata T. 2002. MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform. Nucleic Acids Research 30 (14): 3059–3066.

Kearse M., Moir R., Wilson A., Stones-Havas S., Cheung M., Sturrock S., Buxton S., Cooper A., Markowitz D.C., Thierer T., Ashton B. & Drummond A. 2012. Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics 28 (12): 1647–1649.

Kelaart E.F. 1858. Description of new and little known species of Ceylon nudibranchiate molluscs, and zoophytes. Journal of the Ceylon Branch of the Royal Asiatic Society 3 (9): 84–139. Available from [accessed 17 Apr. 2023].

Klussmann-Kolb A. & Brodie G.D. 1999. Internal storage and production of symbiotic bacteria in the reproductive system of a tropical marine gastropod. Marine Biology 133: 443–447.

Klussmann-Kolb A., Dinapoli A., Kuhn K., Streit B. & Albrecht Ch. 2008. From sea to land and beyond – New insights into the evolution of euthyneuran Gastropoda (Mollusca). BMC Evolutionary Biology 8: 57.

Korshunova T.A., Fletcher K., Picton B., Lundin K., Kashio Sh., Sanamyan N., Sanamyan K., Padula V., Schrödl M. & Martynov A. 2020. The emperor’s Cadlina, hidden diversity and gill cavity evolution: new insights for the taxonomy and phylogeny of dorid nudibranchs (Mollusca: Gastropoda). Zoological Journal of the Linnean Society 20: 1–66.

Leigh J.W. & Bryant D. 2015. Popart: full-feature software for haplotype network construction. Methods in Ecology and Evolution 6: 1110–1116.

Maniei F., Espeland M., Movahedi M. & Wägele H. 2020. Description of a new Peronia species (Gastropoda: Eupulmonata: Onchidiidae) from Iran, Persian Gulf. Zootaxa 4758 (3): 501–531.

Marcus Er. 1961. Opisthobranchs from North Carolina. The Journal of the Elisha Mitchell Scientific Society 77 (2): 141–151.

Marcus Ev. & Marcus Er. 1970. Some gastropods from Madagascar and West Mexico. Malacologia 10 (1): 181–223. Available from [accessed 17 Apr. 2023].

Maroni P. & Wilson N. 2022. Multiple Doris “kerguelenensis” (Nudibranchia) species span the Antarctic Polar Front. Ecology and Evolution 12 (9): e9333.

MolluscaBase eds 2023a. MolluscaBase. Cariopsilla Ortea & Espinosa, 2006. Available from [accessed 25 Aug. 2023].

MolluscaBase eds 2023b. MolluscaBase. Doriopsilla pharpa Er. Marcus, 1961. Accessed from [accessed 5 Sep. 2023].

Mousavipoor Y. 2013. Molecular and Morphological Analysis of some Heterobranchia Species (Mollusca) from Chabahar Coasts. MSc thesis, Marine Biology Department, Chabahar Maritime University. [In Persian.]

Nguyen L.T., Schmidt H.A., von Haeseler A. & Minh B.Q. 2014. IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution 32: 268–274.

Nimbs M.J. & Smith S.D.A. 2021. Genetic evidence confirms that the porostomate nudibranch Dendrodoris gunnamatta Allan, 1932 is a morphotype of Dendrodoris krusensternii (Gray, 1850) (Gastropoda: Nudibranchia). Taxonomy 1: 152–159.

Nithyanandan M. 2012. New and rare nudibranch records from Kuwait, Arabian Gulf (Mollusca: Opisthobranchia). Marine Biodiversity Records 5: 1–7.

Ortea R.J. & Buske Y. 2018. Lista inicial ilustrada de las babosas marinas (Heterobranquios) de la expedición Madibenthos, realizada en 2016 en Martinica (Antillas Menores, mar Caribe). Revista de la Academia Canaria de Ciencias 30: 67–102.

Palumbi S.R., Martin A., Romano S., Owen MacMillan W., Stice L. & Grabowski G. 1991. The Simple Fool’s Guide to PCR. Department of Zoology, University of Hawaii, Honolulu.

Papu A., Undap N., Martinez N.A., Segre M.R., Datang I.G., Kuada R.R., Perin M., Yonow N. & Wägele H. 2020. First study on marine Heterobranchia (Gastropoda, Mollusca) in Bangka Archipelago, North Sulawesi. Indonesia. Diversity 12 (2): 1–72.

Papu A., Bogdanov A., Bara R., Kehraus S., König G.M., Yonow N. & Wägele H. 2022. Phyllidiidae (Nudibranchia, Heterobranchia, Gastropoda): an integrative taxonomic approach including chemical analyses. Organisms Diversity & Evolution 22 (3): 585–629.

Perrone A.S. 2001. A new species of Nudibranchia of the genus Doriopsilla Bergh (Gastropoda, Opisthobranchia) from South Africa. Basteria 65 (1–3): 61–68.

Puillandre N., Lambert A., Brouillet S. & Achaz G. 2011. ABGD: Automatic Barcode Gap Discovery for primary species delimitation. Molecular Ecology 21: 1864–1877.

Pruvot-Fol A. 1951. Étude des Nudibranches de la Méditerranée. Archives de Zoologie expérimentale et générale 88: 1–80.

Rezai H., Mohtarami S.A., Dehghani H., Tavakoli-Kolour P., Bargahi H.R. & Kabiri K. 2016. Nudibranchs from the northern Persian Gulf. Journal of the Persian Gulf (Marine Science) 7 (23): 23–24. Available from [accessed 17 Apr. 2023].

Rüppell E. & Leuckart F.S. 1828–1830. Mollusca. In: Rüppell E. (ed.) Atlas zu der Reise im nördlichen Afrika. Neue wirbellose Thiere des Rothen Meeres: 15–22 [1828]; 23–47 [1830]. H.L. Brönner, Frankfurt. Available from [accessed 17 Apr. 2023].

Stoffels B.E.M.W., van der Meij S.E.T., Hoeksema B.W., van Alphen J., van Alen T., Meyers-Muñoz M.A., de Voogd N.J., Tuti Y. & van der Velde G. 2016. Phylogenetic relationships within the Phyllidiidae (Opisthobranchia, Nudibranchia). ZooKeys 605: 1–35.

Templeton A.R., Crandall K.A. & Sing C.F. 1992. A cladistic analysis of phenotypic associations with haplotypes inferred from restriction endonuclease mapping and DNA sequence data III. Cladogram estimation. Genetics 132: 619–633.

Tibiriçá Y., Pola M. & Cervera J.L. 2017. Astonishing diversity revealed: an annotated and illustrated inventory of Nudipleura (Gastropoda: Heterobranchia) from Mozambique. Zootaxa 4359 (1): 1–133.

Thollesson M. 2020. Increasing fidelity in parsimony analysis of dorid nudibranchs by differential weighting, or a tale of two genes. Molecular Phylogenetics and Evolution 16 (2): 167–172.

Trifinopoulos J., Nguyen L.T., Haeseler A. & Minh B.Q. 2016. W-IQ-TREE: a fast online phylogenetic tool for maximum likelihood analysis. Nucleic Acids Research 44: 232–235.

Valdés Á. 2002. A phylogenetic analysis and systematic revision of the cryptobranch dorids (Mollusca, Nudibranchia, Anthobranchia). Zoological Journal of the Linnean Society 136: 535–636.

Valdés Á. 2003. Preliminary phylogeny of the radula-less dorids (Gastropoda: Opisthobranchia), based on 16S mtDNA sequence data. Journal of Molluscan Studies 69: 75–80.

Valdés Á. & Fahey S.J. 2006. Dorid nudibranchs described by J.E. Gray in M.E. Gray, 1842–1857 (Mollusca: Opisthobranchia). Records of the Western Australian Museum Supplement 69: 95–102.

Valdés Á. & Gosliner T.M. 1999. Phylogenetic systematics of the radula-less dorids (Mollusca: Nudibranchia) with the description of a new genus and a new family. Zoologica Scripta 28: 315–360.

Valdés Á. & Hamann J. 2008. Two new species of Doriopsilla from the tropical Western Atlantic with remarks on Cariopsillidae Ortea & Espinosa, 2005. Veliger 50 (3): 210–218. Available from [accessed 17 Apr. 2023].

Valdés Á. & Ortea J. 1997. Review of the genus Doriopsilla Bergh, 1880 (Gastropoda, Nudibranchia) in the Atlantic Ocean. Veliger 40: 240–254. Available from [accessed 17 Apr. 2023].

Valdés Á., Ortea J., Ávila C. & Ballesteros M. 1996. Review of the genus Dendrodoris Ehrenberg, 1831 (Gastropoda. Nudibranchia) in the Atlantic Ocean. Journal of Molluscan Studies 62: 1–31.

Wägele H., Brodie G. & Klussmann-Kolb A. 1999. Histological investigations of Dendrodoris nigra (Gastropoda, Nudibranchia, Dendrodorididae). Molluscan Research 20: 79–94.

WoRMS Editorial Board 2023. MolluscaBase. Cariopsilla pharpa (Er. Marcus, 1961). Available from [accessed 5 Sep. 2023].

Yonow N. 2008. Sea Slugs of the Red Sea. Pensoft Series Faunistica No. 74, Sofia–Moscow.

Yonow N. 2012. Opisthobranchs from the Western Indian Ocean, with descriptions of two new species and ten new records (Mollusca, Gastropoda). ZooKeys 197: 1–129.

Yonow N. 2015. Sea slugs: unexpected biodiversity and distribution. In: Rasul N.M.A. & Stewart I.C.F. (eds) The Red Sea: 531–550. Springer, Berlin.

How to Cite
Maniei, F., & Wägele, H. (2024). Dendrodorididae (Heterobranchia, Nudibranchia) from Persian Gulf with a description of a new species of Doriopsilla and remarks on the family. European Journal of Taxonomy, 943(1), 179–217.
Research article