Disentangling species of the genus Limacia O.F. Müller, 1781, from southern Africa and Europe using integrative taxonomical methods, with the description of four new species

Alder J, Hancock A (1848) A monograph of the British Nudibranchiate Mollusca, with figures of all the species, part 4. Ray Society, London Altschul SF, Gish W, Miller W, Myers EW, Lipman DJ (1990) Basic local alignment search tool. J Mol Biol 215(3):403–410. https://doi.org/10.1016/S0022-2836(05)80360-2 Ari J (2014) Orange-clubbed Sea Slug (Limacia clavigera). iNaturalist. https://www.inaturalist.org/observations/9493589. Accessed 10 May 2019 Ari J (2016) Orange-clubbed Sea Slug (Limacia clavigera). iNaturalist. https://www.inaturalist.org/observations/9657487. Accessed 10 May 2019 Baba K (1937) Opisthobranchia of Japan (II). Journal of the Department of Agriculture, Kyushu Imperial University 5(7):289–344 Bacon CD, Silvestro D, Jaramillo C, Smith BT, Chakrabarty P, Antonelli A (2015) Biological evidence supports an early and complex emergence of the isthmus of Panama. P Natl Acad Sci USA 112(19):6110–6115. https://doi.org/10.1073/pnas.1423853112 Ballesteros M, Madrenas E, Pontes M (2012-2020) "Limacia clavigera" in OPK-Opistobranquis. http://opistobranquis.info/en/Exy8r. Accessed 10 May 2019 Barnard KH (1927) South African nudibranch Mollusca, with descriptions of new species, and a note on some specimens from Tristan d'Acunha. Annals of the South African Museum 25(1):171–215 Benson DA, Cavanaugh M, Clark K, Karsch-Mizrachi I, Lipman DJ, Ostell J, Sayers EW (2017) GenBank. Nucleic Acids Res 45(D1):D37–D42. https://doi.org/10.1093/nar/gkw1070 Bergh LSR (1884) Report on the Nudibranchiata dredged by H.M.S. Challanger during the years 1873–1876. In: Report on the scientific results of the voyage of H.M.S. Challenger during the years 1873–76 under the command of Captain George S. Nares and the late Captain Frank Tourle Thomson. Zoology Vol. X. Neil and Company, Edinburgh Bergh LSR (1907) The Opisthobranchiata of South Africa. T S Afr Philos Soc 17(1):1–144 Bertsch H, Ferreira AJ (1974) Four new species of nudibranchs from tropical west America. Cuatro especies nuevas de nudibranquios de la América tropical occidental. Veliger 16(4):343–353 Bickford D, Lohman DJ, Sodhi NS, Ng PK, Meier R, Winker K, Ingram KK, Das I (2007) Cryptic species as a window on diversity and conservation. Trends Ecol Evol 22(3):148–155. https://doi.org/10.1016/j.tree.2006.11.004 Bolton JJ, Anderson RJ, Smit AJ, Rothman MD (2012) South African kelp moving eastwards: the discovery of Ecklonia maxima (Osbeck) Papenfuss at De Hoop Nature Reserve on the south coast of South Africa. Afr J Mar Sci 34(1):147–151. https://doi.org/10.2989/1814232X.2012.675125 Boonzaaier MK (2017) Diversity and zoogeography of South African Bryozoa. Dissertation, University of the Western Cape, South Africa Bourjea J, Lapegue S, Gagnevin L, Broderick D, Mortimer JA, Ciccione S, Roos D, Taquet C, Grizel H (2007) Phylogeography of the green turtle, Chelonia mydas, in the Southwest Indian Ocean. Mol Ecol 16(1):175–186. https://doi.org/10.1111/j.1365-294X.2006.03122.x Bowen BW, Grant WS (1997) Phylogeography of the sardines (Sardinops spp.): assessing biogeographic models and population histories in temperate upwelling zones. Evolution 51(5):1601–1610. https://doi.org/10.1111/j.1558-5646.1997.tb01483.x Bowen BW, Gaither MR, DiBattista JD, Iacchei M, Andrews KR, Grant WS, Toonen RJ, Briggs JC (2016) Comparative phylogeography of the ocean planet. P Natl Acad Sci USA 113(29):7962–7969. https://doi.org/10.1073/pnas.1602404113 Buyukbaykal F (2003) Limacia clavigera from Turkey. Sea Slug Forum. Australian Museum, Sydney. http://www.seaslugforum.net/find/10776. Accessed 10 May 2019 Caballer Gutiérrez M, Almón BA, Pérez JP (2016) The sea slug genus Limacia Müller, 1781 (Mollusca: Gastropoda: Heterobranchia) in Europe. Cah Biol Mar 57:35–42 Calado GJP, Silva JP (2012) Lesmas do Mar do Algarve: Guia de moluscos opistobrânquios da costa sul de Portugal. SubNauta, pp:978–989 Cantraine FJ (1835) Diagnoses ou descriptiones succinctes de quelques espèces nouvelles de mollusques. Bulletin de l'Académie Royale des Sciences et Belles-lettres de Bruxelles 2(11):380–401 Carmona L, Bhave V, Salunkhe R, Pola M, Gosliner TM, Cervera JL (2014) Systematic review of Anteaeolidiella (Mollusca, Nudibranchia, Aeolidiidae) based on morphological and molecular data, with a description of three new species. Zool J Linnean Soc 171(1):108–132. https://doi.org/10.1111/zoj.12129 Churchill CK, Valdés Á, Foighil DÓ (2014) Molecular and morphological systematics of neustonic nudibranchs (Mollusca: Gastropoda: Glaucidae: Glaucus), with descriptions of three new cryptic species. Invertebr Syst 28(2):174–195. https://doi.org/10.1071/IS13038 Cole M, Ranwashe F (2018) FBIP:ELM: malacology collections. South African National Biodiversity Institute Occurrence dataset https://doi.org/10.15468/uyagu4 Accessed (via GBIF.org) 10 May 2019 Coppard SE, Lessios HA (2017) Phylogeography of the sand dollar genus Encope: implications regarding the Central American Isthmus and rates of molecular evolution Sci Rep 7:11520. https://doi.org/10.1038/s41598-017-11875-w Cuvier GLCF (1817) Le règne animal distribué d'après son organisation : pour servir de base a l'histoire naturelle des animaux et d'introduction a l'anatomie comparée. Déterville, Paris Debelius H, Kuiter RH (2007) Nudibranchs of the world. Ikan-Unterwasserarchiv, Frankfurt Felsenstein J (1978) Cases in which parsimony or compatibility methods will be positively misleading. Syst Zool 27(4):401–410. https://doi.org/10.2307/2412923 Floeter SR, Rocha LA, Robertson DR, Joyeux JC, Smith-Vaniz WF, Wirtz P, Edwards AJ, Barreiros JP, Ferreira CE, Gasparini JL, Brito A (2008) Atlantic reef fish biogeography and evolution. J Biogeogr 35(1):22–47. https://doi.org/10.1111/j.1365-2699.2007.01790.x Florence WK, Gibbons MJ, Hayward PJ (2007) Taxonomy of shallow-water Bryozoa from the west coast of South Africa. African Natural History 3(1):1–58 Gaither MR, Rocha LA (2013) Origins of species richness in the Indo-Malay-Philippine biodiversity hotspot: evidence for the centre of overlap hypothesis. J Biogeogr 40(9):1638–1648. https://doi.org/10.1111/jbi.12126 García-Gómez JC (2002) Paradigmas de una fauna insólita. Los moluscos opistobranquios del Estrecho de Gibraltar. Instituto de Estudios Campogibraltareños, Algeciras, Cádiz GBIF Secretariat (2017a) Limacia O.F. Müller, 1781. GBIF Backbone Taxonomy. https://www.gbif.org/species/2291771. Accessed 5 Sep 2018 GBIF Secretariat (2017b) Limacia clavigera (O.F. Müller, 1776). GBIF Backbone Taxonomy. https://www.gbif.org/species/5190033. Accessed 5 Sep 2018 Geller J, Meyer C, Parker M, Hawk H (2013) Redesign of PCR primers for mitochondrial cytochrome c oxidase subunit I for marine invertebrates and application in all-taxa biotic surveys. Mol Ecol Resour 13(5):851–861. https://doi.org/10.1111/1755-0998.12138 Ghanimi H, Schrödl M, Goddard JH, Ballesteros M, Gosliner TM, Buske Y, Valdés Á (2020) Stargazing under the sea: molecular and morphological data reveal a constellation of species in the Berthella stellata (Risso, 1826) species complex (Mollusca, Heterobranchia, Pleurobranchidae). Mar Biodivers 50(1):11. https://doi.org/10.1007/s12526-019-01027-w Glon H (2020) Orange-clubbed Sea Slug (Limacia clavigera). iNaturalist. https://www.inaturalist.org/observations/28462590. Accessed 20 Dec 2019 Goddard JH, Treneman N, Prestholdt T, Hoover C, Green B, Pence WE, Mason DE, Dobry P, Sones JL, Sanford E, Agarwal R (2018) Heterobranch sea slug range shifts in the northeast Pacific Ocean associated with the 2015–16 El Niño. Proc Calif Acad Sci 4(65):107–131 Golestani H, Crocetta F, Padula V, Camacho-García Y, Langeneck J, Poursanidis D, Pola M, Yokeş MB, Cervera JL, Jung DW, Gosliner TM (2019) The little Aplysia coming of age: from one species to a complex of species complexes in Aplysia parvula (Mollusca: Gastropoda: Heterobranchia). Zool J Linnean Soc 187(2):279–330. https://doi.org/10.1093/zoolinnean/zlz028 Gosliner TM (1987) Nudibranchs of southern Africa: a guide to opisthobranch molluscs of southern Africa. Sea Challengers, Monterey, California Gosliner TM, Behrens DW, Valdés Á (2008) Indo-Pacific nudibranchs and sea slugs: a field guide to the world’s most diverse fauna. Sea Challengers Natural History Books, California Academy of Sciences, San Francisco Gosliner TM, Behrens DW, Valdés Á (2015) Nudibranch & sea slug identification: Indo-Pacific, 1st edn. New World Publications, Jacksonville Gosliner TM, Behrens DW, Valdés Á (2018) Nudibranch & sea slug identification: Indo-Pacific, 2nd edn. New World Publications, Jacksonville Grant WS, Bowen BW (2006) Living in a tilted world: climate change and geography limit speciation in Old World anchovies (Engraulis; Engraulidae). Biol J Linn Soc 88(4):673–689 Hajibabaei M, Singer GA, Hebert PD, Hickey DA (2007) DNA barcoding: how it complements taxonomy, molecular phylogenetics and population genetics. Trends Genet 23(4):167–172. https://doi.org/10.1016/j.tig.2007.02.001 Hallas JM, Chichvarkhin A, Gosliner TM (2017) Aligning evidence: concerns regarding multiple sequence alignments in estimating the phylogeny of the Nudibranchia suborder Doridina. R Soc Open Sci 4(10):171095. https://doi.org/10.1098/rsos.171095 Hamon N, Sepulchre P, Lefebvre V, Ramstein G (2013) The role of eastern Tethys seaway closure in the Middle Miocene Climatic Transition (ca. 14 Ma). Clim Past 9(6):2687–2702. https://doi.org/10.5194/cp-9-2687-2013 Hendy MD, Penny D (1989) A framework for the quantitative study of evolutionary trees. Syst Zool 38(4):297–309. https://doi.org/10.2307/2992396 Henriques R, Potts WM, Santos CV, Sauer WH, Shaw PW (2014) Population connectivity and phylogeography of a coastal fish, Atractoscion aequidens (Sciaenidae), across the Benguela current region: evidence of an ancient vicariant event. PLoS One 9(2):e87907. https://doi.org/10.1371/journal.pone.0087907 Hering M (1841) Diagnoses algarum novarum a cl. Dr Ferdinand Krauss in Africa australi lectarum. Ann Mag Nat Hist 8(49):90–92 Hoareau TB, Boissin E, Paulay G, Bruggemann JH (2013) The Southwestern Indian Ocean as a potential marine evolutionary hotspot: perspectives from comparative phylogeography of reef brittle-stars. J Biogeogr 40(11):2167–2179. https://doi.org/10.1111/jbi.12155 Horst D (2008) Re: Limacia clavigera from French Mediterranean. Sea Slug Forum. Australian Museum, Sydney. http://www.seaslugforum.net/find/21456. Accessed 10 May 2019 Hubbs CL (1952) Antitropical distribution of fishes and other organisms. Symposium on problems of bipolarity and of pantemperate faunas Proc Seventh Pac Sci Congr (Pac Sci Assoc) 3:324–329 Huelsenbeck JP, Ronquist F (2001) MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics 17(8):754–755. https://doi.org/10.1093/bioinformatics/17.8.754 Hutchings L, van der Lingen CD, Shannon LJ (2009) The Benguela Current: an ecosystem of four components. Prog Oceanogr 83(1–4):15–32. https://doi.org/10.1016/j.pocean.2009.07.046 iNaturalist (2020a) Observations of Limacia in South Africa. https://www.inaturalist.org/observations?place_id=6986&taxon_id=47766. Accessed 10 Jan 2020 iNaturalist (2020b) Observations of Limacia in Europe. https://www.inaturalist.org/taxa/47766-Limacia/browse_photos?layout=grid&place_id=97391. Accessed 10 Jan 2020 Jacobs DK, Haney TA, Louie KD (2004) Genes, diversity, and geologic process on the Pacific coast. Annu Rev Earth Planet Sci 32:601–652. https://doi.org/10.1146/annurev.earth.32.092203.122436 Johnston G (1834) Magazine of natural history and journal of zoology, botany, mineralogy, geology and meteorology, Vol. VII, London Johnston G (1838) Annals of natural history, vol I. R. and J.E, Taylor, London Jones G (2008) A field guide to the marine animals of the cape peninsula. Southern Underwater Research Group, Cape Town Kearse M, Moir R, Wilson A, Stones-Havas S, Cheung M, Sturrock S, Buxton S, Cooper A, Markowitz S, Duran C, Thierer T (2012) Geneious basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics 28(12):1647–1649. https://doi.org/10.1093/bioinformatics/bts199 Kimura M (1980) A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences. J Mol Evol 16(2):111–120. https://doi.org/10.1007/BF01731581 Knowlton N (1993) Sibling species in the sea. Annu Rev Ecol Syst 24(1):189–216. https://doi.org/10.1146/annurev.es.24.110193.001201 Kumar S, Stecher G, Tamura K (2016) MEGA7: molecular evolutionary genetics analysis version 7.0 for bigger datasets. Mol Biol Evol 33(7):1870–1874. https://doi.org/10.1093/molbev/msw054 Larkin MA, Blackshields G, Brown NP, Chenna R, McGettigan PA, McWilliam H, Valentin F, Wallace IM, Wilm A, Lopez R, Thompson JD, Gibson TJ, Higgins DG (2007) Clustal W and Clustal X version 2.0. Bioinformatics 23(21):2947–2948. https://doi.org/10.1093/bioinformatics/btm404 Lima E (2014) in: Casassovici A, Brosens D (2019) Diveboard - Scuba diving citizen science observations. https://www.gbif.org/occurrence/1020055551. Accessed 5 Sep 2018 Lindberg DR (1991) Marine biotic interchange between the northern and southern hemispheres. Paleobiology 17(3):308–324. https://doi.org/10.1017/S0094837300010629 Linnaeus C (1767) Systema naturae per regna tria naturae: secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Ed. 12. 1., Regnum Animale. 1 & 2. Holmiae, Laurentii Salvii. Holmiae [Stockholm], Laurentii Salvii. pp 533–1327 Lockhart P, Steel M (2005) A tale of two processes. Syst Biol 54(6):948–951. https://doi.org/10.1080/10635150500234682 Losos JB, Arnold SJ, Bejerano G, Brodie ED III, Hibbett D, Hoekstra HE, Mindell DP, Monteiro A, Moritz C, Orr HA, Petrov DA, Renner SS, Ricklefs RE, Soltis PS, Turner TL (2013) Evolutionary biology for the 21st century. PLoS Biol 11(11):e1001466. https://doi.org/10.1371/journal.pbio.1001466 Ludt WB, Rocha LA (2015) Shifting seas: the impacts of Pleistocene Sea-level fluctuations on the evolution of tropical marine taxa. J Biogeogr 42(1):25–38. https://doi.org/10.1111/jbi.12416 Ludt WB, Rocha LA, Erdmann MV, Chakrabarty P (2015) Skipping across the tropics: the evolutionary history of sawtail surgeonfishes (Acanthuridae: Prionurus). Mol Phylogenet Evol 84:166–172. https://doi.org/10.1016/j.ympev.2014.12.017 Lundin KG (2019) Record details for SWEMA758-15. Barcode of life data System v4. http://boldsystems.org/index.php/Public_RecordView?processid=SWEMA758-15. Accessed 1 May 2019 MacFarland FM (1905) A preliminary account of the Dorididae of Monterey Bay, California. Proc Biol Soc Wash 18:35–54 Macnae W (1957) The families Polyceridae and Goniodorididae (Mollusca, Nudibranchiata) in southern Africa. Transactions of the Royal Society of South Africa 35(4):341–372 Maggs CA, Castilho R, Foltz D, Henzler C, Jolly MT, Kelly J, Olsen J, Perez KE, Stam W, Väinölä R, Viard F (2008) Evaluating signatures of glacial refugia for North Atlantic benthic marine taxa. Ecology 89(sp11):S108–S122. https://doi.org/10.1890/08-0257.1 Malaquias MAE, Valdés Á, Alexander J, Schrödl M, Hooker Y, Padula V, Gosliner TM, Bottomley M, Camacho Y, Breslau E, Vital XG (2018) Molecular and morphological systematics of Dolabrifera gray, 1847 (Mollusca: Gastropoda: Heterobranchia: Aplysiomorpha). Zool J Linnean Soc 184(1):31–65. https://doi.org/10.1093/zoolinnean/zlx099 Marcus E (1964) A new species of Polycera (Nudibranchia) from California. Nautilus 77(4):128–131 Marlow JR, Lange CB, Wefer G, Rosell-Melé A (2000) Upwelling intensification as part of the Pliocene-Pleistocene climate transition. Science 290(5500):2288–2291. https://doi.org/10.1126/science.290.5500.2288 Martínez-Chacón M (2018) Nudibranquios de la Isla de Tarifa. Pequeñas joyas del estrecho de Gibraltar. Consejería de Medio Ambiente y Ordenación del Territorio, Junta de Andalucía. Sevilla Mayer AG (1918) Biographical memoir of William Stimpson, 1832–1872: presented to the Academy at the Annual Meeting, 1917. National Academy of sciences Mayes (2008) Oman Oct 2008. Flickr, Brian Mayes https://www.flickr.com/photos/brianmayes/3053688644/in/album-72157608962836782/. Accessed 10 May 2019 Miller BJ, von der Heyden S, Gibbons MJ (2012) Significant population genetic structuring of the holoplanktic scyphozoan Pelagia noctiluca in the Atlantic Ocean. Afr J Mar Sci 34(3):425–430. https://doi.org/10.2989/1814232X.2012.726646 MolluscaBase (2019) Limacia O.F. Müller, 1781. World register of marine species. http://wwwmarinespeciesorg/aphiaphp?p=taxdetails&id=138368 on 2019-06-06. Accessed 6 June 2019 Montagu G (1813). Descriptions of several new or rare animals, principally marine, discovered on the south coast of Devonshire. T Linn Soc London 11(1):1–26. https://doi.org/10.1111/j.1096-3642.1813.tb00035.x Müller OF (1776) Zoologiae Danicae prodromus: seu Animalium Daniae et Norvegiae indigenarum characteres, nomina, et synonyma imprimis popularium. Typis Hallageriis. https://doi.org/10.5962/bhl.title.13268 Müller OF (1781) Zoologia Danica eller Danmarks og Norges sieldne og ubekiendte dyrs historie. Hallager, Copenhagen Müller OF (1788) Zoologia danica: sev Animalivm Daniae et Norvegiae rariorvm ac minvs notorvm descriptiones et historia. Typis N. Mölleri avlae regiae typographie et filii. Copenhagen. https://doi.org/10.5962/bhl.title.49477 Muths D, Tessier E, Bourjea J (2014) Genetic structure of the reef grouper Epinephelus merra in the West Indian Ocean appears congruent with biogeographic and oceanographic boundaries. Mar Ecol 36(3):447–461. https://doi.org/10.1111/maec.12153 Nylander JAA (2004) MrModeltest v2.3 software. Program distributed by the author. Evolutionary Biology Center, Uppsala University, Sweden O’Dea A, Lessios HA, Coates AG, Eytan RI, Restrepo-Moreno SA, Cione AL, Collins LS, De Queiroz A, Farris DW, Norris RD, Stallard RF (2016) Formation of the isthmus of Panama. Sci Adv 2(8):e1600883. https://doi.org/10.1126/sciadv.1600883 Odhner NH (1941) New polycerid nudibranchiate Mollusca and remarks on this family. Elanders boktryck, Gothenburg Ores S (2006) Limacia clavigera from Brittany, France. Sea Slug Forum. Australian Museum, Sydney. http://www.seaslugforum.net/find/15508. Accessed 10 May 2019 Ortea J, Quero A, Rodriguez G, Valdés Á (1989) Study of Limacia clavigera (Muller, 1776) (Mollusca: Nudibranchia), with notes about its geographical distribution and on the validity of the genus Laila MacFarland, 1905. Revista de Biologia de la Universidad de Oviedo (Spain) Ortea J, Moro L, Benhamú C, Bacallado JJ, Ocaña O (2018) Babosas marinas de la region de Ceuta. Fundación Museo del Mar de Ceuta. Grafisur Tarifa SL, Cádiz Palomar G, Pola M, Garcia-Vazquez E (2014) First molecular phylogeny of the subfamily Polycerinae (Mollusca, Nudibranchia, Polyceridae). Helgoland Mar Res 68(1):143–153. https://doi.org/10.1007/s10152-013-0374-z Papenfuss GF (1940) Notes on South African marine algae. I Bot Notiser 1940:200–226 Poddubetskaia M (2002) Limacia clavigera from French Brittany. Sea Slug Forum. Australian Museum, Sydney. http://www.seaslugforum.net/find/7831. Accessed 10 May 2019 Pola M, Gosliner TM (2010) The first molecular phylogeny of cladobranchian opisthobranchs (Mollusca, Gastropoda, Nudibranchia). Mol Phylogenet Evol 56(3):931–941. https://doi.org/10.1016/j.ympev.2010.05.003 Pola M, Cervera JL, Gosliner TM (2006) Description of two new phanerobranch nembrothid species (Nudibranchia: Polyceridae: Doridacea). J Mar Biol Assoc UK 86(2):403–409. https://doi.org/10.1017/S0025315406013269 Pola M, Cervera JL, Gosliner TM (2007) Phylogenetic relationships of Nembrothinae (Mollusca: Doridacea: Polyceridae) inferred from morphology and mitochondrial DNA. Mol Phylogenet Evol 43(3):726–742. https://doi.org/10.1016/j.ympev.2007.02.003 Pola M, Camacho-Garcia YE, Gosliner TM (2012) Molecular data illuminate cryptic nudibranch species: the evolution of the Scyllaeidae (Nudibranchia: Dendronotina) with a revision of Notobryon. Zool J Linnean Soc 165(2):311–336. https://doi.org/10.1111/j.1096-3642.2012.00816.x Prkić J, Petani A, Iglić D, Lanča L (2018) Opisthobranchs of the Adriatic Sea. Photographic atlas and list of Croatian species. Diving Club St Roko, Bibinje Puillandre N, Lambert A, Brouillet S, Achaz G (2012) ABGD, automatic barcode gap discovery for primary species delimitation. Mol Ecol 21(8):1864–1877. https://doi.org/10.1111/j.1365-294X.2011.05239.x Randall JE (1981) Examples of anti-tropical and anti-equatorial distribution of Indo-West-Pacific fishes. Pac Sci 35(3):197–209 Rambaut A (2016) Figtree v1.4.3. Computer program distributed by the author, University of Edinburgh. http://tree.bio.ed.ac.uk/software/figtree/ Rambaut A, Drummond AJ, Xie D, Baele G, Suchard MA (2018) Posterior summarisation in Bayesian phylogenetics using Tracer 1.7. Syst Biol. syy032. https://doi.org/10.1093/sysbio/syy032 Ratnasingham S, Hebert PD (2007) BOLD: the barcode of life data system (http://www. barcodinglife. org). Mol Ecol Notes 7(3):355–364. https://doi.org/10.1111/j.1471-8286.2007.01678.x Rudman WB (1999) Limacia clavigera (Muller, 1776). Sea Slug Forum. Australian Museum, Sydney. http://www.seaslugforum.net/find/limaclav. Accessed 10 May 2019 Rudman WB (2003) Radula of Limacia ornata. Sea Slug Forum. Australian Museum, Sydney. http://www.seaslugforum.net/find/8920. Accessed 10 May 2019 Sánchez-Tocino L (2006) Opistobranquios. Fauna Andaluza. pp 115-184. En Tinaut A, Pascual F (Coordinadores). Proyecto Andalucía. Naturaleza. Volumen XIV (Zoología II). Publicaciones Comunitarias, SL, Sevilla, Spain Schmekel L, Portmann A (1982) Opisthobranchia des Mittelmeeres. Springer Science & Business Media, Berlin, Heidelberg Schmittner A, Sarnthein N, Kinkel H, Bartoli G, Bickert T, Crucifix M, Crudeli D, Groeneveld J, Kösters F, Mikolajewicz U, Millo C (2004) Global impact of the Panamanian seaway closure. Eos, Transactions American Geophysical Union 85(49):526–526. https://doi.org/10.1029/2004EO490010 Sellick B, Gosliner TM, van Heerden I (2020) Nudibranchs and sea slugs of the Aliwal Shoal. Pietermaritzburg, South Africa Shipman C, Gosliner T (2015) Molecular and morphological systematics of Doto Oken, 1851 (Gastropoda: Heterobranchia), with descriptions of five new species and a new genus. Zootaxa 3973(1):57–101. https://doi.org/10.11646/zootaxa.3973.1.2 Silva JP (2007) Limacia clavigera and possible egg mass. Sea Slug Forum. Australian Museum, Sydney. http://www.seaslugforum.net/find/20643. Accessed 10 May 2019 Skauge C (2007) Limacia clavigera mating. Sea Slug Forum. Australian Museum, Sydney. http://www.seaslugforum.net/find/21097. Accessed 10 May 2019 Spalding MD, Fox HE, Allen GR, Davidson N, Ferdaña ZA, Finlayson MA, Halpern BS, Jorge MA, Lombana AL, Lourie SA, Martin KD (2007) Marine ecoregions of the world: a bioregionalization of coastal and shelf areas. BioScience 57(7):573–583. https://doi.org/10.1641/B570707 Stadler T (2009) On incomplete sampling under birth–death models and connections to the sampling-based coalescent. J Theor Biol 261(1):58–66. https://doi.org/10.1016/j.jtbi.2009.07.018 Stamatakis A (2014) RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics 30(9):1312–1313. https://doi.org/10.1093/bioinformatics/btu033 Stamatakis A (2015) Using RAxML to infer phylogenies. Curr Protoc Bioinformatics 51(1):6–14. https://doi.org/10.1002/0471250953.bi0614s51 Stimpson W (1855) Description of some new marine invertebrates. Proc Acad Natl Sci Phila 7(10):385–395 Struck TH, Feder JL, Bendiksby M, Birkeland S, Cerca J, Gusarov VI, Kistenich S, Larsson KH, Liow LH, Nowak MD, Stedje B (2018) Finding evolutionary processes hidden in cryptic species. Trends Ecol Evol 33(3):153–163. https://doi.org/10.1016/j.tree.2017.11.007 Suchard MA, Lemey P, Baele G, Ayres DL, Drummond AJ, Rambaut A (2018) Bayesian phylogenetic and phylodynamic data integration using BEAST 1.10 virus Evolution 4, vey016. https://doi.org/10.1093/ve/vey016 Teske PR, von der Heyden S, McQuaid CD, Barker NP (2011) A review of marine phylogeography in southern Africa. S Afr J Sci 107(5–6):43–53. https://doi.org/10.4102/sajs.v107i5/6.514 Thollesson M (2000) Increasing fidelity in parsimony analysis of dorid nudibranchs by differential weighting, or a tale of two genes. Mol Phylogenet Evol 16(2):161–172. https://doi.org/10.1006/mpev.2000.0789 Thompson W (1840) Annals of natural history, Vol. V. R. and J.E. Taylor Thompson TE, Brown GH (1976) British opisthobranch molluscs. Published for the Linnean Society of London by Academic Press Thompson TE, Brown GH (1984) Biology of opisthobranch molluscs, vol II. Ray Society, London Tibiriçá Y, Pola M, Cervera JL (2017) Astonishing diversity revealed: an annotated and illustrated inventory of Nudipleura (Gastropoda: Heterobranchia) from Mozambique. Zootaxa 4359(1):1–133. https://doi.org/10.11646/zootaxa.4359.1.1 Toms JA, Compton JS, Smale M, von der Heyden S (2014) Variation in palaeo-shorelines explains contemporary population genetic patterns of rocky shore species. Biol Lett 10(6):20140330. https://doi.org/10.1098/rsbl.2014.0330 Trivedi S, Aloufi AA, Ansari AA, Ghosh SK (2016) Role of DNA barcoding in marine biodiversity assessment and conservation: an update. Saudi J Biol Sci 23(2):161–171. https://doi.org/10.1016/j.sjbs.2015.01.001 Uribe RA, Sepúlveda F, Goddard JH, Valdés Á (2017) Integrative systematics of the genus Limacia O.F. Müller, 1781 (Gastropoda, Heterobranchia, Nudibranchia, Polyceridae) in the Eastern Pacific. Mar Biodivers 48(4):1815–1832. https://doi.org/10.1007/s12526-017-0676-5 Vallès YV, Valdés Á, Ortea JE (2000) On the phanerobranch dorids of Angola (Mollusca, Nudibranchia): a crossroads of temperate and tropical species. Zoosystema 22(1):15–32 van Bragt PH (1999) Re: Limacia clavigera from the Netherlands. Sea Slug Forum, Australian Museum, Sydney http://www.seaslugforum.net/find/1106. Accessed 10 May 2019 Vermeij GJ (1992) Trans-equatorial connections between biotas in the temperate eastern Atlantic. Mar Biol 112(2):343–348. https://doi.org/10.1007/BF00702481 von der Heyden S (2017) Making evolutionary history count: biodiversity planning for coral reef fishes and the conservation of evolutionary processes. Coral Reefs 36(1):183–194. https://doi.org/10.1007/s00338-016-1512-2 Wägele H, Willan RC (2000) Phylogeny of the Nudibranchia. Zool J Linnean Soc 130(1):83–181. https://doi.org/10.1111/j.1096-3642.2000.tb02196.x Weaver AJ (1990) Ocean currents and climate. Nature 347(6292):432–432 Wells FE, Bryce CW (1993) Sea slugs and their relatives of western Australia. Western Australian Museum, Perth Winnepenninckx B, Backeljau T, De Wachter R (1993) Extraction of high molecular weight DNA from molluscs. Trends Genet 9(12):407. https://doi.org/10.1016/0168-9525(93)90102-n Wirtz P (1998) Opisthobranch molluscs from the Azores. Vita Marina 45(1–2):1–16 Womersley HBS (1996) The marine benthic flora of southern Australia - part IIIB - Gracilariales, Rhodymeniales, Corallinales and Bonnemaisoniales. Canberra & Adelaide: Australian Biological Resources Study & the State Herbarium of South Australia 5:1–392 Xiong B, Kocher TD (1991) Comparison of mitochondrial DNA sequences of seven morphospecies of black flies (Diptera: Simuliidae). Genome 34(2):306–311. https://doi.org/10.1139/g91-050 Zhang J, Kapli P, Pavlidis P, Stamatakis A (2013) A general species delimitation method with applications to phylogenetic placements. Bioinformatics 29(22):2869–2876. https://doi.org/10.1093/bioinformatics/btt499 Zsilavecz (2007) Nudibranchs of the cape peninsula and False Bay. Southern Underwater Research Group, Cape Town