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Article
Published: 2020-05-15

There are three species of Chrysaora (Scyphozoa: Discomedusae) in the Benguela upwelling ecosystem, not two

Department of Biodiversity and Conservation Biology, University of the Western Cape, Private Bag X17, Bellville 7535, South Africa.
Department of Biodiversity and Conservation Biology, University of the Western Cape, Private Bag X17, Bellville 7535, South Africa.
Department of Biodiversity and Conservation Biology, University of the Western Cape, Private Bag X17, Bellville 7535, South Africa.
Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Rua do Matão trav. 14, n. 101, São Paulo, SP, 05508-090, BRAZIL.
Noblis ESI, 112 Industrial Park Boulevard, Warner Robins, United States, GA 31088.
Department of Biodiversity and Conservation Biology, University of the Western Cape, Private Bag X17, Bellville 7535, South Africa. 7National Marine and Information Research Centre (NatMIRC), Ministry of Fisheries and Marine Resources, P.O.Box 912, Swakopmund, Namibia.
Department of Biodiversity and Conservation Biology, University of the Western Cape, Private Bag X17, Bellville 7535, South Africa.
Coelenterata Agulhas Bank Benguela upwelling region Namibia New species Pelagiidae Scyphozoa Taxonomy

Abstract

Chrysaora (Pèron & Lesueur 1810) is the most diverse genus within Discomedusae, and 15 valid species are currently recognised, with many others not formally described. Since Chrysaora fulgida (Reynaud 1830) was first recognised as occurring off the south west (SW) coast off South Africa, the species has been variously synonymised with Chrysaora hysoscella (Linnaeus 1767) and Chrysaora africana (Vanhöffen 1902). Using DNA evidence alongside multivariate tools to analyse quantitative morphometric and meristic data, as well as information from the cnidome, we unambiguously separate C. fulgida from C. hysoscella; we resurrect C. africana as a valid species and recognise a new species, Chrysaora agulhensis sp. nov. Full descriptions of C. fulgida, C. africana and C. agulhensis sp. nov. are provided. The species have different geographical patterns of distribution around the region, with restricted areas of overlap: C. agulhensis sp. nov. is found along the southern coast of South Africa and over the Agulhas Bank, C. fulgida extends from Cape Point in South Africa to southern Angola, and C. africana can be found from southern Namibia northwards to the Gulf of Guinea. The species can be readily separated in the field by a combination of tentacle/lappet number and shape, colour patterns and the form of the oral arms.

 

References

  1. Abboud, S.S, Gómez Daglio, L. & Dawson, M.N. (2018) A global estimate of genetic and geographic differentiation in macromedusae—implications for identifying the causes of jellyfish blooms. Marine Ecology Progress Series, 591, 199­–216.

    https://doi.org/10.3354/meps12521

    Agassiz, L. (1862) Contributions to the Natural History of the United States of America. Vol. IV. Pt. III. Discophorae. Pt. IV. Hydroidae. Pt. V. Homologies of the Radiata. Little Brown Trubner, Boston, London, 380 pp.

    Akaike, H. (1973) Information theory and an extension of the maximum likelihood principle. In: Petrov, P.N. & Csaki, F. (Eds.), Second International Symposium on Information Theory. Adad. Kiado, Budapest, pp. 267–281.

    Anderson, M.J., Gorley, R.N. & Clarke, K.R. (2008) PERMANOVA+ for PRIMER: Guide to Software and Statistical Methods. PRIMER-E, Plymouth, 214 pp.

    Awad, A.A., Griffiths, C.L. & Turpie, J.K. (2002) Distribution of South African marine invertebrates applied to the selection of priority conservation areas. Diversity & Distribution, 8, 129–145.

    https://doi.org/10.1046/j.1472-4642.2002.00132.x

    Bayha, K.M., Collins, A.G. & Gaffney, P.M. (2017) Multigene phylogeny of the scyphozoan jellyfish family Pelagiidae reveals that the common U.S. Atlantic sea nettle comprises two distinct species (Chrysaora quinquecirrha and C. chesapeakei) PeerJ, 5, e3863.

    https://doi.org/10.7717/peerj.3863

    Bayha, K.M., Dawson, M.N., Collins, A.G., Barbeitos, M.S. & Haddock, S.H.D. (2010) Evolutionary relationships among scyphozoan jellyfish families based on complete taxon sampling and phylogenetic analysis of 18S and 28S ribosomal DNA. Integrative and Comparative Biology, 50 (3), 436–455.

    https://doi.org/10.1093/icb/icq074

    Bologna, P., Gaynor, J.J., Meredith, R., Restaino, D. & Barry, C. (2018) Stochastic event alters gelatinous zooplankton community structure: impacts of Hurricane Sandy in a Mid-Atlantic estuary. Marine Ecology Progress Series, 591, 217−227.

    https://doi.org/10.3354/meps12262

    Brierley, A.S., Axelsen, B.E., Buecher, E., Sparks, C., Boyer, H. & Gibbons, M.J. (2001) Acoustic observations of jellyfish in the Namibian Benguela. Marine Ecology Progress Series, 210, 55–66.

    https://doi.org/10.3354/meps210055

    Brierley, A.S., Axelsen, B.E., Boyer, D.C., Lynam, C.P., Didcock, C.A., Boyer, H.J., Sparks, C.A.J., Purcell, J.E. & Gibbons, M.J. (2004) Single-target echo detections of jellyfish. ICES Journal of Marine Science, 61, 383–393.

    https://doi.org/10.1016/j.icesjms.2003.12.008

    Brodeur, R.D., Link, J.S., Smith, B.E., Ford, M., Kobayashi, D. & Jones, T.T. (2016) Ecological and economic consequences of ignoring jellyfish:a plea for increased monitoring of ecosystems. Fisheries, 41, 630−637.

    https://doi.org/10.1080/03632415.2016.1232964

    Buecher, E., Sparks, C., Brierley, A., Boyer, H. & Gibbons, M.J. (2001) Biometry and size distribution of Chrysaora hysoscella (Cnidaria, Scyphozoa) and Aequorea aequorea (Cnidaria, Hydrozoa) off Namibia with some notes on their parasite Hyperia medusarum. Journal of Plankton Research, 23, 1073–1080.

    https://doi.org/10.1093/plankt/23.10.1073

    Carlgren, O. (1940) A contribution to the knowledge of the structure and distribution of the cnidae in Anthozoa. Lunds Universitet´s Årskrift, 36, 1–62.

    Darriba, D., Taboada, GL., Doallo, R. & Posada, D. (2012) jModelTest 2: more models, new heuristics and parallel computing. Nature Methods, 9, 772.

    https://doi.org/10.1038/nmeth.2109

    Dawson, M.N. (2003) Macro-morphological variation among cryptic species of the moon jellyfish, Aurelia (Cnidaria: Scyphozoa). Marine Biology, 143, 369–379.

    https://doi.org/10.1007/s00227-003-1070-3

    Dawson, M.N. (2005) Morphological variation and systematics in the Scyphozoa: Mastigias (Rhizostomeae, Mastigiidae)—a golden unstandard? Hydrobiologia, 537, 185–206.

    https://doi.org/10.1007/s10750-004-2840-8

    Dawson, M.N. & Jacobs, D.K. (2001) Molecular evidence for cryptic species of Aurelia aurita (Cnidaria, Scyphozoa). Biological Bulletin, 200, 92–96.

    https://doi.org/10.2307/1543089

    de Lafontaine, Y. & Leggett, W.C. (1989) Changes in size and weight of hydromedusae during formalin preservation. Bulletin of Marine Science, 44, 1129–1137.

    Desor, E. (1848) No title. Meeting of the November 1, 1848. Proceedings of the Boston Society of Natural History, 3 (1848–1851), 73–77. [Desor part, pp. 75–76.]

    https://doi.org/10.1080/03745485809494477

    Dwivedi, B. & Gadagkar, S.R. (2009) Phylogenetic inference under varying proportions of indel-induced alignment gaps. BMC Evolutionary Biology, 9, 211.

    https://doi.org/10.1186/1471-2148-9-211

    Fearon, J.J., Boyd, A.J. & Schülein, F.H. (1992) Views on the biomass and distribution of Chrysaora hysoscella (Linné) and Aequorea aequorea (Forskål, 1775) off Namibia 1982–1989. Scientia Marina, 56, 75–85.

    Feigenbaum, D.L. & Kelly, M. (1984) Changes in the lower Chesapeake Bay food chain in the presence of the sea nettle Chrysaora quinquecirrha (Scyphomedusae). Marine Ecology Progress Series, 19, 39–47

    https://doi.org/10.3354/meps019039

    Felsenstein, J. (1985) Confidence limits on phylogenies: An approach using the bootstrap. Evolution, 39, 783–791

    https://doi.org/10.1111/j.1558-5646.1985.tb00420.x

    Flynn, B.A. & Gibbons, M.J. (2007) A note on the diet and feeding of Chrysaora hysoscella in Walvis Bay Lagoon, Namibia, during September 2003. African Journal of Marine Science, 29, 303–307.

    https://doi.org/10.2989/ajms.2007.29.2.15.197

    Folmer, O., Black, M., Hoeh, W., Lutz, R. & Vrijenhoek, R. (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3, 294–299.

    Gegenbaur, C. (1856) Versuch eines Systemes der Medusen, mit Beschreibung neuer oder wenig gekannter Formen; zugleich ein Beitrag zur Kenntnis der Fauna des Mittelmeeres. Zeitschrift für Wissenschaftliche Zoologie, Leipzig, 8, 202–273.

    Gershwin, L. & Collins, A.G. (2002) A preliminary phylogeny of Pelagiidae (Cnidaria, Scyphozoa), with new observations of Chrysaora colorata comb. nov. Journal of Natural History, 36, 127–148.

    https://doi.org/10.1080/00222930010003819

    Gómez-Daglio, L. (2016) Systematics and phylogeny of shallow water jellyfish (Scyphozoa, Discomedusae in the Tropical Eastern Pacific. PhD thesis. Accessible from: https://escholarship.org/uc/item/03s3r0qf (accessed 23 March 2013)

    Gómez-Daglio, L. & Dawson, M.N. (2017) Species richness of jellyfishes (Scyphozoa:Discomedusae) in the Tropical Eastern Pacific: missed taxa, molecules, and morphology match in a biodiversity hotspot. Invertebrate Systematics, 31, 635−663.

    https://doi.org/10.1071/IS16055

    Griffiths, C.L., Robinson, T.B., Lange, L. & Mead, A. (2010) Marine biodiversity in South Africa–state of knowledge, spatial patterns and threats. PLoS ONE, 5 (8), e123008.

    https://doi.org/10.1371/journal.pone.0012008

    Hall, T. (2005) BioEdit, Biological sequence alignment editor for Win95/98/NT/2K/XP. Available from: www.mbio.ncsu.edu/BioEdit/bioedit.html. (accessed 23 March 2013)

    Haeckel, E. (1880) 2: System der Acraspeden. In: Das System der Medusen. I. Gustav Fischer, Jena, pp. 361–672.

    Holland, B., Dawson, M., Crow, G. & Hofmann, D. (2004) Global phylogeography of Cassiopea (Scyphozoa: Rhizostomeae): molecular evidence for cryptic species and multiple invasions of the Hawaiian Islands. Marine Biology, 145, 1119–1128.

    https://doi.org/10.1007/s00227-004-1409-4

    Hutchings, L., Barange, M., Bloomer, S.F., Boyd, A.J., Crawford, R.J.M., Huggett, J.A., Kerstan, M., Korrûbel, J.L., de Oliveira, J.A.A., Painting, S.J., Richardson, A.J., Schülein, F.H., van der Lingen, C.D. & Verheye, H.M. (1998) Multiple factors affecting South African anchovy recruitment in the spawning, transport and nursery areas. South African Journal of Marine Science, 19 (1), 211–225.

    https://doi.org/10.2989/025776198784126908

    Hutchings, L., Van der Lingen, C.D., Shannon, L.J., Crawford, R.J.M., Verheye, H.M.S., Bartholomae, C.H., Van der Plas, A.K., Louw, D., Kreiner, A., Ostrowski, M., Fidel, Q., Barlow, R.G., Lamont, T., Coetzee, J., Shillington, F., Veitch, J., Currie, J.C. & Monteiro, P.M.S. (2009) The Benguela Current: an ecosystem of four components. Progress in Oceanography, 83 (1–4), 15–32.

    https://doi.org/10.1016/j.pocean.2009.07.046

    Kramp, P.L. (1955) The medusae of the tropical west coast of Africa. Atlantide Reports, 3, 239–324.

    Kramp, P.L. (1961) Synopsis of the medusae of the world. Journal of Marine Biological Association of the United Kingdom, 40, 1–469.
    https://doi.org/10.1017/s0025315400007347

    Kumar, S., Stecher, G., Li, M., Knyaz, C. & Tamura, K. (2018) MEGA X: Molecular Evolutionary Genetics Analysis across computing platforms. Molecular Biology and Evolution, 35, 1547–1549.

    https://doi.org/10.1093/molbev/msy096

    Lesson, R.P. (1843) Histoire naturelle des Zoophytes, Acalèphes. Librairie Encyclopédique de Roret, Paris, 596 pp.

    https://doi.org/10.5962/bhl.title.4799

    Linnaeus, C. (1767) Systema naturae per regna tria naturae: secundum classes, ordines, genera, species cum characteribus, differentiis, sinonimis, locis. Editio duodecima, reformata. Tomus I. Pars II. Laurentii Salvii, Holmiae, 550 pp.

    https://doi.org/10.5962/bhl.title.158187

    Lucas, C.H. (2009) Biochemical composition of the mesopelagic coronate jellyfish Periphylla periphylla from the Gulf of Mexico. Journal of the Marine Biological Association of the United Kingdom, 89, 77–81

    https://doi.org/10.1017/S0025315408002804

    Lucas, C.H., Graham, W.M. & Widmer, C. (2012) Jellyfish life histories: role of polyps in forming and maintaining scypho-medusa populations. Advances in Marine Biology, 63, 133−196.

    https://doi.org/10.1016/B978-0-12-394282-1.00003-X

    Lutjeharms, J.R.E. & Cooper, J. (1996) Inter-basin leakage through Agulhas Current filaments. Deep-Sea Research, 43, 213–238

    . https://doi.org/10.1016/0967-0637(96)00002-7

    Lynam, C.P., Gibbons, M.J., Axelsen, B.A., Sparks, C.A.J., Coetzee, J., Heywood, B.G. & Brierley, A.S. (2006) Jellyfish overtake fish in a heavily fished ecosystem. Current Biology, 16, 492–493.

    https://doi.org/10.1016/j.cub.2006.06.018

    Lynch, M. & Crease, T.J. (1990). The analysis of population survey data on DNA sequence variation. Molecular Biology and Evolution, 7, 377–394.

    https://doi.org/10.1093/oxfordjournals.molbev.a040607

    Maposa, S. (2005) Jellyfish plague hobbles Koeberg. Available from: https://www.iol.co.za/news/south-africa/jellyfish-plague-hobbles-koeberg-240851 (accessed 21 September 2015)

    Mariottini, G.L., Giacco, E. & Pane, L. (2008) The mauve stinger Pelagia noctiluca (Forsskal, 1775). Distribution, ecology, toxicity and epidemiology of stings. A review. Marine Drugs, 6, 496–513.

    https://doi.org/10.3390/md20080025

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

    https://doi.org/10.1016/B978-0-12-512150-7.50008-6

    Mayer, A.G. (1910) Volume III: Schyphomedusae. In: Mayer, A.G., Medusae of the world. Carnegie Institute, Washington, pp. 499–735.

    https://doi.org/10.5962/bhl.title.159245

    McClain, C.R., Balk, M.A., Benfield, M.C., Branch, T.A., Chen, C., Cosgrove, J., Dove, A.D.M., Gaskins, L.C., Helm, R.R., Hochberg, F.G., Lee, F.B., Marshall, A., McMurray, S.E., Schanche, C., Stone, S.N. & Thaler, A.D. (2015) Sizing ocean giants: patterns of intraspecific size variation in marine megafauna. PeerJ, 3, e715

    https://doi.org/10.7717/peerj.715

    Medlin, L.K., Elwood, H.J., Stickel, S. & Sogin, M.L. (1988) The characterization of enzymatically amplified eukaryotic 16S-like rRNA-coding regions. Gene, 71, 491–499.

    https://doi.org/10.1016/0378-1119(88)90066-2

    Mejía-Sánchez, N. & Marques, A.C. (2013) Getting information from ethanol preserved nematocysts of the venomous cubomedusa Chiropsalmus quadrumanus: a simple technique to facilitate the study of nematocysts. Latin American Journal of Aquatic Research, 41 (1), 166–169.

    https://doi.org/103856/vol41-issue1-fulltext-14

    Mello, B. (2018) Estimating TimeTrees with MEGA and the TimeTree Resource. Molecular Biology and Evolution, 35 (9), 2334–2342

    https://doi.org/10.1093/molbev/msy133

    Mianzan, H.W. & Cornelius, P.F.S. (1999) Cubomedusae and Scyphomedusae. In: Boltovskoy, D. (Ed.), South Atlantic Zooplankton. Vol. 1. Backhuys Publishers, Leiden, pp. 513–559.

    Miller, G.A. & Chapman, J.P. (2001) Misunderstanding analysis of covariance. Journal of Abnormal Psychology, 110 (1), 40–48.

    https://doi.org/10.1037/0021-843X.110.1.40

    Mills, C.E. (2001) Jellyfish blooms: are populations increasing globally in response to ocean conditions? Hydrobiologia, 451, 55–68.

    https://doi.org/10.1007/978-94-010-0722-1_6

    Morandini, A.C. & Marques, A.C. (2010) Revision of the genus Chrysaora Péron & Lesuer, 1810 (Cnidaria: Scyphozoa). Zootaxa, 2464 (1), 1–97.

    https://doi.org/10.11646/zootaxa.2464.1.1

    Neethling, S. (2010) Re-descriptions of some South African scyphozoa: out with the old and in with the new. MSc thesis University of the Western Cape, University of the Western Cape, 172 pp. [unpublished notes]

    Nei, M. (1987) Molecular Evolutionary Genetics. Columbia University Press, New York, 512 pp.

    https://doi.org/10.7312/nei-92038

    Nei, M. & Kumar, S. (2000) Molecular Evolution and Phylogenetics. Oxford University Press, New York, 333 pp.

    Nylander, J.A.A. (2004) MrModeltest. Version 2. Program Distributed by the Author. Evolutionary Biology Centre, Uppsala University, Uppsala, Sweden. [software]

    Ogden, T. & Rosenberg, M. (2007) How should gaps be treated in parsimony? A comparison of approaches using simulation. Molecular phylogenetics and evolution, 42, 817–826.

    https://10.1016/j.ympev.2006.07.021

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

    https://doi.org/10.3989/scimar.2000.64s131

    Pagès, F., Gili, J.M. & Bouillon, J. (1992) Medusae (Hydrozoa, Scyphozoa, Cubozoa) of the Benguela Current (southeastern Atlantic). In: Pagès, F., Gili, J.M. & Bouillon, J. (Eds.), Planktonic Cnidarians of the Benguela Current. Scientia Marina, 56 (Supplement 1), pp. 1–64.

    Palomares, M.L.D. & Pauly, D. (2009) The growth of jellyfishes. Hydrobiologia, 616, 11–21.

    https://doi.org/10.1007/s10750-008-9582-y

    Papenfuss, E.J. (1936) The utility of the nematocysts in the classification of certain Scyphomedusae. I. Cyanea capillata, Cyanea palmstruchii, Dactylometra quinquecirrha, Dactylometra quinquecirrha var. chesapeakei, and Chrysaora hysoscella. Acta Universitatis Lundensis, Nova Series, 31 (11), 19–26.

    Peach, M.B. & Pitt, K.A. (2005) Morphology of the nematocysts of the medusae of two scyphozoans, Catostylus mosaicus and Phyllorhzia punctata (Rhizostomeae): implications for capture of prey. Invertebrate biology, 124 (2), 98–108.

    https://doi.org/10.1111/j.1744-7410.2005.00012.x

    Péron, F. & Lesueur, C.A. (1810) Tableau des caractères génériques et spécifiques de toutes les espèces de Méduses connues jusqu‟à ce jour. Annales du Muséum National d’Histoire Naturelle, Paris, 14, 325–366.

    Purcell, J.E., Uye, S. & Lo, W. (2007) Anthropogenic causes of jellyfish blooms and their direct consequences for humans: a review. Marine Ecology Progress Series, 350, 153–174.

    https://doi.org/10.3354/meps07093

    Quinn, G. & Keough, M. (2002) Experimental Design and Data Analysis for Biologists. Cambridge University Press, Cambridge, 537 pp.

    https://doi.org/10.1017/cbo9780511806384

    Ramhaut, A. (2009) FigTree. Version 1.4.1 Available from: http://tree.bio.ed.ac.uk/software/figtree/ (accessed 8 May 2013)

    Ramhaut, A. & Drummond, A.J. (2007) Tracer. Version 1.5. Available from: http://beast.bio.ed.ac.uk/Tracer/ (accessed 8 May 2013)

    Ranson, G. (1949) Résultats scientifiques des croisières du navire école belge “Mercator” IV. II Méduses. Mémoires du Institut Royal des Sciences Naturelles de Belgique, Série 2, 33, 121–158.

    Reynaud, A.A.M. (1830) Medusa (Cyanea) caliparea; Medusa (Rhyzostoma) fulgida. In: Lesson, R.P. (Ed.), Centurie Zoologique, ou choix d’animaux rares, nouveaux ou imparfaitement connus. F.G. Levrault, Paris, pp. 67–68, 79–80.

    Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574.

    https://doi.org/10.1093/bioinformatics/btg180

    Sambrook, J. & Russell, D.W. (2001) Molecular Cloning: A Laboratory Manual. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, New York, 34 pp.

    Sink, K.J., Gibbons, M.J., Laird, M.C. & Atkinson, L.J. (2017) Phylum Cnidaria In: Atkinson, L.J. & Sink, K.J. (Eds.), Field Guide to the Offshore Marine Invertebrates of South Africa, Malachite Marketing and Media, Pretoria, pp. 65–115.

    Sparks, C., Buecher, E., Brierley, A.S., Boyer, H., Axelsen, B.E. & Gibbons, M.J. (2001) Observations on the distribution and relative abundance of the scyphomedusan Chrysaora hysoscella (Linné, 1766) and the hydrozoan Aequorea aequorea (Forskål, 1775) in the northern Benguela ecosystem. Hydrobiologia, 451, 275–286.

    https://doi.org/10.1023/A:1011829516239

    Stiasny, G. (1934) Die rhizostomeen-sammlung des British Museum (Natural History) in London. Zoologische Mededeelingen, 14, 137–178.

    Stiasny, G. (1939) Über Dactylometra fulgida (Reynaud) von der Walfischbai. Zoologischer Anzeiger, 126, 172–185.

    Straehler-Pohl, I. & Jarms, G. (2010) Identification key for young ephyrae: a first step for early detection of jellyfish blooms. Hydrobiologia, 645, 3–21.

    https://doi.org/10.1007/s10750-010-0226-7

    Suchman, C.L., Daly, E.A., Keister, J.E., Peterson, W.T. & Brodeur, R.D. (2008) Feeding patterns and predation potential of scyphomedusae in a highly productive upwelling region. Marine Ecology Progress Series, 358, 161–172.

    https://doi.org/10.3354/meps07313

    Swofford, D. (2003) PAUP—Phylogenetic Analysis Using Parsimony (* and Other Methods). Version 4.0b10. Sinauer Associates, Sunderland, Massachusetts. [software]

    Tajima, F. (1983) Evolutionary relationship of DNA sequences in finite populations. Genetics, 105, 437–460.

    Tamura, K., Battistuzzi. F.U., Billing-Ross, P., Murillo, O., Filipski, A. & Kumar, S. (2012) Estimating Divergence Times in Large Molecular Phylogenies. Proceedings of the National Academy of Sciences, 109, 19333–19338.

    https://doi.org/10.1073/pnas.1213199109

    Thiel, H. (1966) The evolution of the Scyphozoa, a review. In: Rees, W.J. (Ed.), Cnidaria and their Evolution. Academic Press, London, pp. 77–117.

    Thibault-Botha, D. & Bowen, T. (2004) Impact of formalin preservation on Pleurobrachia bachei (Ctenophora). Journal of Experimental Marine Biology and Ecology, 303, 11–17.

    https://doi.org/10.1016/j.jembe.2003.10.017

    van der Lingen, C.D., Shannon, L.J., Cury, P., Kreiner, A., Moloney, C.L., Roux, J.-P. & Vaz- Velho, F. (2006) Resource and Ecosystem Variability, Including Regime Shifts, in the Benguela Current System. In: Shannon, V., Hempel, G., Malanotte-Rizzoli, P., Moloney, C. & Woods, J. (Eds.), Large Marine Ecosystems. Vol. 14. Elsevier, pp. 146–184.

    https://doi.org/10.1016/s1570-0461(06)80013-3

    Vanhöffen, E. (1888) Untersuchungen über semäostome und rhizostome Medusen. Bibliotheca Zoologica, 1, 5–52.

    https://doi.org/10.5962/bhl.title.38091

    Vanhöffen, E. (1902) Die Acraspeden Medusen der deutschen Tiefsee-expedition 1898–1899. Wissenschaftliche Ergebnisse der deutschen Tiefsee-expedition auf dem dampfer Valdivia 1898–1899, 3, 3–52.

    https://doi.org/10.5962/bhl.title.1404

    Venter, G.E. (1988) Occurrence of jellyfish on the west coast off south west Africa/Namibia, in Long-term data series relating to southern Africa’s renewable natural resources. South African National Science Progress Reports, 157, 56–61.

    Vousden, D., Stapley, J.R., Ngoile, M.A.K., Sauer, W. & Scott, L. (2012) Climate change and variability of the Agulhas and Somali Current Large Marine Ecosystem in relation to socioeconomics and governance. Chapter 5 in: Frontline observations on climate change and sustainability of Large Marine Ecosystems, 17, 81–96.

    Wallace, D.M. (1987) Large- and small-scale phenol extractions. Methods in Enzymology, 152, 33–41.

    https://doi.org/10.1016/0076-6879(87)52007-9

    Widmer, C.L. (2005) Effects of temperature on growth of north-east Pacific moon jellyfish ephyrae, Aurelia labiata (Cnidaria: Scyphozoa). Journal of the Marine Biological Association of the United Kingdom, 85, 569–573.

    https://doi.org/10.1017/S0025315405011495

    Zar, J.H. (1999) Biostatistical Analysis. 4th Edition. Dorling Kindersley, New Delhi, 663 pp.