Skip to main content
Log in

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

  • Original Paper
  • Published:
Marine Biodiversity Aims and scope Submit manuscript


Doriopsilla areolata is a nudibranch sea slug with a broad geographic range, spanning from mainland Spain (southwestern Europe) to Angola (southwestern Africa) in the eastern Atlantic, and is also found in the Caribbean. This species was divided into three subspecies: D. areolata areolata (Europe to Cape Verde), D. areolata albolineata (Ghana to Angola), and D. areolata nigrolineata (Caribbean), each with a discrete range. This paper uses phylogenetic and population genetics tools to determine whether the subspecies scheme is supported by molecular data. Additionally, specimens of Doriopsilla miniata from South Africa, a species that is hypothetically closely related to D. areolata, is also included in the analysis, primarily because of the morphological similarities between D. areolata and D. miniata. Results from AMOVA, Bayesian, and pairwise analyses revealed that the molecular data do not support the proposed subspecies scheme and suggest that South African populations identified as D. miniata belong to D. areolata.

This is a preview of subscription content, log in via an institution to check access.

Access this article

Price excludes VAT (USA)
Tax calculation will be finalised during checkout.

Instant access to the full article PDF.

Fig. 1
Fig. 2

Similar content being viewed by others


  • Akaike H (1974) A new look at the statistical model identifications. IEEE Trans Autom Control 19:716–723. doi:10.1109/TAC.1974.1100705

    Article  Google Scholar 

  • 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. Trans Zool Soc Lond 5:113–147

    Article  Google Scholar 

  • Alfaro ME, Zoller S, Lutzoni F (2003) Bayes or Bootstrap? A simulation study comparing the performance of Bayesian Markov Chain Monte Carlo sampling and bootstrapping in assessing phylogenetic confidence. Mol Biol Evol 20:255–266. doi:10.1093/molbev/msg028

    Article  PubMed  CAS  Google Scholar 

  • Bergh LSR (1880) Die Doriopsen des Mittelmeeres. Jahrb Dtsch Malakozool Ges 7:297–328

    Google Scholar 

  • Burn R (2006) A checklist and bibliography of the Opisthobranchia (Mollusca: Gastropoda) of Victoria and the Bass Strait area, south-eastern Australia. Mus Vic Sci Rep 10:1–42

    Google Scholar 

  • Calado G, Soares C, Neves R (2012) Reproduction in two sympatric Iberian nudibranch species (Gastropoda: Opisthobranchia) with contrasting development types. Iberus 30:103–110

    Google Scholar 

  • Clement M, Posada D, Crandall K (2000) TCS: a computer program to estimate gene genealogies. Mol Ecol 9:1657–1660. doi:10.1046/j.1365-294x.2000.01020.x

    Article  PubMed  CAS  Google Scholar 

  • Colgan DJ, McLauchlan A, Wilson GDF, Livingston SP, Edgecombe GD, Macaranas J, Cassis G, Gray MR (1998) Histone H3 and U2 snRNA DNA sequences and arthropod molecular evolution. Aust J Zool 46:419–437

    Article  Google Scholar 

  • Drummond AJ, Kearse M, Heled J, Moir R, Thierer T, Ashton B, Wilson A, Stones-Havas S (2009) Geneious v4.8. Accessed 03 March 2010

  • Duran S, Palacín C, Becerro MA, Turon X, Giribet G (2004) Genetic diversity and population structure of the commercially harvested sea urchin Paracentrotus lividus (Echinodermata, Echinoidea). Mol Ecol. doi:10.11111/j1365-294X.2004.02338.x

  • Edmunds M (1968) Opistobranchiate Mollusca from Ghana. Proc Malacol Soc Lond 38:83–100

    Google Scholar 

  • Excoffier L, Lischer HEL (2010) Arlequin suite ver 3.5: a new series of programs to perform population genetics analyses under Linux and Windows. Mol Ecol Resour 10:564–567. doi:10.1111/j.1755-0998.2010.02847.x

    Article  PubMed  Google Scholar 

  • Farris JS, Källersjö M, Kluge AG, Bult C (1994) Testing significance of incongruence. Cladistics 10:315–319

    Article  Google Scholar 

  • Gosliner T (1987) Nudibranchs of Southern Africa. A guide to Opisthobranch Molluscs of Southern Africa. Sea Challengers, Monterey

    Google Scholar 

  • Huelsenbeck JP, Ronquist F (2001) MrBayes: Bayesian inference of phylogenetic trees. Bioinformatics 17:754–755. doi:10.1093/bioinformatics/17.8.754

    Article  PubMed  CAS  Google Scholar 

  • Lessios HA, Kessing BD, Pearse JS (2001) Population structure and speciation in tropical seas: global phylogeography of the sea urchin Diadema. Evol 55:955–975

    Article  CAS  Google Scholar 

  • Meyer KB (1977) Dorid nudibranchs of the Caribbean coast of the Panama Canal Zone. Bull Mar Sci Gulf Caribb 27:299–307

    Google Scholar 

  • Mickevich MF, Farris JS (1981) The implications of congruence in Menidia. Syst Zool 30:351–370

    Article  Google Scholar 

  • Nylander JAA (2004) MrModeltest v2. Evolutionary Biology Centre, Uppsala University Accessed 22 May 2008

  • Ornelas-Gatdula E, Dupont A, Valdés A (2011) The tail tells the tale: taxonomy and biogeography of some Atlantic Chelidonura (Gastropoda: Cephalaspidea: Aglajidae) inferred from nuclear and mitochondrial gene data. Zool J Linn Soc 163:1077–1095. doi:10.1111/j.1096-3642.2011.00749.x

    Article  Google Scholar 

  • Palumbi SR (1996) Nucleic acids II: The polymerase chain reaction. In: Hillis DM, Moritz C, Mable BK (eds) Molecular Systematics. Sinauer, Sunderland, pp 205–247

    Google Scholar 

  • Pérez-Losada M, Nolte MJ, Crandall KA, Shaw PW (2007) Testing hypotheses of population structuring in the Northeast Atlantic Ocean and Mediterranean Sea using the common cuttlefish Sepia officinalis. Mol Ecol 16:2667–2679. doi:10.11111/j.1365-294X.2007.03333.x

    Article  PubMed  Google Scholar 

  • Rose RA (1985) The spawn and development of twenty-nine New South Wales opisthobranchs (Mollusca: Gastropoda). Proc Linn Soc N S W 108:23–36

    Google Scholar 

  • Swofford DL (2002) PAUP*: Phylogenetic analysis using parsimony (*and other methods). Ver. 4. Sinauer, Sunderland

    Google Scholar 

  • Uliana E, Lange CB, Wefer G (2002) Evidence for Congo River freshwater load in Late Quaternary sediments of ODP Site 1077 (5ºS, 10ºE). Palaeogeogr Palaeocl 187:137–150. doi:10.1016/j.bbr.2011.03.031

    Article  Google Scholar 

  • Valdés A, Ortea J (1997) Review of the Genus Doriopsilla Bergh, 1880 (Gastropoda, Nudibranchia) in the Atlantic Ocean. Veliger 40:240–254

    Google Scholar 

  • Valdés A, Hamann J (2008) Two new species of Doriopsilla from the tropical Western Atlantic with remarks on Cariopsillidae Ortea & Espinosa, 2005. Veliger 50:210–218

    Google Scholar 

  • Xavier R, Zenboudji S, Lima FP, Harris DJ, Santos AM, Branco M (2011) Phylogeography of the marine isopod Stenosoma nadejda (Rezig, 1989) in North African Atlantic and western Mediterranean coasts reveals complex differentiation patterns and a new species. Biol J Linn Soc 104:419–431

    Article  Google Scholar 

  • Xia X, Lemey P (2009) Assessing substitution saturation with DAMBE. In: Lemey P, Salemi M, Vandamme A-M (eds) The phylogenetic handbook: A practical approach to DNA and protein phylogeny, 2nd edn. Cambridge University Press, Cambridge, pp 615–630

    Chapter  Google Scholar 

  • Xia X, Xie Z (2001) DAMBE: software package for data analysis in molecular biology and evolution. J Hered 92:371–373. doi:10.1093/jhered/92.4.371

    Article  PubMed  CAS  Google Scholar 

  • Xia X, Xie Z, Salemi M, Chen L, Wang Y (2003) An index of substitution saturation and its application. Mol Phyl Evol 26:1–7. doi:10.1016/S1055-7903(02)00326-3

    Article  CAS  Google Scholar 

Download references


We are extremely grateful to Lindsey Groves from the Natural History Museum of Los Angeles County and Rafael Araujo from the Museo Nacional de Ciencias Naturales in Madrid, Spain (MNCN) for their assistance with access to their collections. Terry Gosliner, Rebecca Johnson and Elizabeth Kools kindly gave us access to the California Academy of Sciences collection. Two anonymous reviewers provided comments that greatly improved the quality of this paper.

Author information

Authors and Affiliations


Corresponding author

Correspondence to Ángel Valdés.

Rights and permissions

Reprints and permissions

About this article

Cite this article

Goodheart, J., Valdés, Á. 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. Mar Biodiv 43, 113–120 (2013).

Download citation

  • Received:

  • Revised:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI: