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Taxonomic transfer of the red algae Chrysymenia enteromorpha and C. wrightii to the genus Botryocladia (Rhodymeniaceae, Rhodymeniales)



Genera in the Rhodymeniaceae Harvey that have a hollow thallus lacking diaphragms have been placed in Chrysymenia J. Agardh 1842 (including Gloiosaccion Harvey 1859), Botryocladia (J. Agardh) Kylin 1931, Irvinea Guiry in Saunders et al. 1999, and Cresia C. Lozada-Troche, D.L. Ballantine & H. Ruíz 2010. Chrysymenia has traditionally been defined by a lack of internal rhizoids and with the only solid portion of the thallus limited to the stipe. Botryocladia has been differentiated from Chrysymenia by the presence of larger, solid axes. Our results suggest that on the basis of vegetative characters alone, the true Chrysymenia clade contains specimens that exclusively produce gland cells directly on unmodified medullary cells, whereas Botryocladia and Cresia may or may not cut off gland cells from modified gland-supporting medullary cells. On the basis of comparative rbcL and SSU sequence analyses and morphological data, the hollow, gelatinous species Chrysymenia enteromorpha Harvey from the Gulf of Mexico and Caribbean Sea, and Chrysymenia wrightii Harvey described from the northwest Pacific Ocean are newly transferred to the genus Botryocladia as Botryocladia enteromorpha (Harvey) comb. nov. and B. wrightii (Harvey) comb. nov., respectively.


algae, biodiversity, Botryocladia, Chrysymenia, Gulf of Mexico, marine, new combination, morphology, phylogeny, Puerto Rico, rbcL, Rhodophyta, Rhodymeniaceae, seaweeds, SSU rDNA

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Abbott, I.A. & Littler, M.M. (1969) Some Rhodymeniales from Hawaii. Phycologia 8: 165–169.

Afonso-Carillo, J. & Sobrino, C. (2003) Vegetative and reproductive morphology of Botryocladia botryoides, B. occidentalis and B. canariensis sp. nov. (Rhodymeniaceae, Rhodophyta) from the Canary Islands. Phycologia 42: 138–150.

Afonso-Carrillo, J., Rodríguez-Prieto, J.C., Boisset, F., Sobrino, C., Tittley, I. & Neto, A.I. (2006) Botryocladia chiajeana and Botryocladia macaronesica sp. nov. (Rhodymeniaceae, Rhodophyta) from the Mediterranean and the Eastern Atlantic, and a discussion on the closely related genus Irvinea. Phycologia 45: 277–292.

Agardh, J.G. (1842) Algae Maris Mediterranei et Adriatici. Apud Fortin, Masson et Cie, Paris, 264 pp.

Ballantine, D.L. & Aponte, N.E. (2002) A checklist of the benthic marine algae known to Puerto Rico, second revision. Constancea 83, online continuation of California Publications in Botany (1902–2002). Available from: (accessed 1 October 2017)

Ballantine, D.L., Ruiz, H. & Lozada-Troche, C. (2010) Chamaebotrys prolifera sp. nov. (Rhodymeniaceae, Rhodophyta) from Puerto Rico, Caribbean Sea, based on morphology and small subunit rDNA sequences. Phycological Research 58: 69–77.

Bárbara, I., Lee, S., Peña, V., Díaz, P., Cremades, J., Oak, J.H. & Choi, H. (2008) Chrysymenia wrightii (Rhodymeniales, Rhodophyta) – a new non-native species for the European Atlantic coast. Aquatic Invasions 3: 367–75.

Ben Maïz, N., Boudouresque, C.F., Riouall, R. & Lauret, M. (1987) Flore algale de l’Etang de Thau (France, Méditerranée): sur la présence d’une Rhodymeniale d’origine japonaise, Chrysymenia wrightii (Rhodophyta). Botanica Marina 30: 357–364.

Børgesen, F. (1920) The marine algae of the Danish West Indies. Part 3. Rhodophyceae (6). Dansk Botanisk Arkiv 3: 369–504.

Brodie, J. & Guiry, M.D. (1988) Life history of Boryocladia ardreana sp. nov. (Rhodymeniales, Rhodophyta) from Portugal. Phycologia 27: 109–130.

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

Dawes, C.J. & Mathieson, A.C. (2008) The Seaweeds of Florida. University Press of Florida. Gainesville, 592 pp.

Felder, D.L., Thoma, B.P., Schmidt, W.E., Sauvage, T., Self-Krayesky, S.L., Chistoserdov, A., Bracken-Grissom, H.D. & Fredericq, S. (2014) Seaweeds and decapod crustaceans on Gulf deep banks after the Macondo Oil Spill. BioScience 64: 808–19.

Feldmann, G. (1945) Révision du genre Botryocladia Kylin (Rhodophycées Rhodyméniacées). Bulletin de la Société d’Histoire Naturelle de l’Afrique du Nord 35: 49–61.

Feldmann, G. & Bodard, M. (1965) Une nouvelle espèce de Botryocladia des côtes du Sénégal. Bulletin de l’Institut Océanographique de Monaco 65: 1–14.

Felsenstein, J. (1985) Phylogenies and the comparative method. The American Naturalist 125: 1015.

Fredericq S., Cho, T.O., Earle, S.A., Gurgel, C.F.D., Krayesky, M., Mateo Cid, L.E., Mendoza Gonzáles, A.C., Norris, J.N. & Suárez, A.M. (2009) Seaweeds of the Gulf of Mexico. In: Felder, D.L. & Camp, D.K. (Eds.) Gulf of Mexico: Its Origins, Waters, and Biota. I. Biodiversity. Texas A&M Univ. Press, pp. 187–259.

Fredericq, S., Arakaki, N., Camacho, O., Gabriel, D., Krayesky, D., Self-Krayesky, S., Rees, G., Richards, J., Sauvage, T., Venera-Ponton, D. & Schmidt, W.E. (2014) A dynamic approach to the study of rhodoliths: a case study for the northwestern Gulf of Mexico. Cryptogamie Algologie 35: 77–98.

Gavio, B. & Fredericq, S. (2003) Botryocladia caraibica sp. nov. (Rhodymeniales, Rhodophyta), a new species from the Caribbean. Cryptogamie Algologie 24: 93–106.

Gavio, B. & Fredericq, S. (2005) New species and new records of offshore members of the Rhodymeniales (Rhodophyta) in the Northern Gulf of Mexico. Gulf of Mexico Science 1: 58–83.

Guiry, M.D. & Guiry, G.M. (2017) Algaebase. Worldwide electronic publication, National University of Ireland, Galway. Available from: (accessed January 2017)

Harvey, W.H. (1853) Nereis Boreali Americana Part II. Rhodospermeae. Smithsonian Contributions to Knowledge 5 (5): 1–258.

Harvey, W.H. (1857) Australian Algae. Exsiccata nos. 1–511, 511 bis, 512–601 (with printed names).

Harvey, W.H. (1859a) Phycologia Australica: or, a history of Australian seaweeds; comprising coloured figures and descriptions of the more characteristic marine algae of New South Wales, Victoria, Tasmania, South Australia, and Western Australia, and a synopsis of all known Australian algae. Vol. 2. Lovell Reeve & Co., London, Pp. [i]–viii, Pls. LXI–CXX (with text).

Harvey, W.H. (1859b) Algae. In: Hooker, J.D. (Ed.) The Botany of the Antarctic Voyage of H.M. Discovery Ships Erebus and Terror, in the Years 1839–1843, Under the Command of Captain Sir James Clark Ross [...] Part III: Flora Tasmaniae. Monocotyledones and Acotyledones, Vol. II. Lovell Reeve, London, pp. 282–343.

Harvey, W.H. (1860) Characters of new algae, chiefly from Japan and adjacent regions, collected by Charles Wright in the North Pacific Exploring Expedition under Captain James Rodgers. Proceedings of the American Academy of Arts and Sciences 4: 327–335.

Haroun, R.J., Gil-Rodríguez, M.C., Díaz de Castro, J. & Prud’homme van Reine, W.F. (2002) A checklist of the marine plants from the Canary Islands (Central eastern Atlantic Ocean). Botanica Marina 45: 139–169.

Hasegawa, M., Kishino, H. & Yano, T. (1985) Dating of the human-ape splitting by a molecular clock of mitochondrial DNA. Journal of Molecular Evolution 22: 160–174.

Hommersand, M.H. & Fredericq, S. (1990) Sexual reproduction and cystocarp development. Pp. 305–345. In: Biology of the Red Algae. Cole, K.M. & Sheath, R.G. [Eds]. Cambridge University Press.

Hommersand, M.H., Fredericq, S. & Cabioch, J. (1992) Developmental morphology of Gigartina pistillata (Gigartinaceae, Rhodophyta). Phycologia 31: 300–325.

Huelsenbeck, J.P. & Ronquist, F. (2001) MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics 17: 754–755.

Kylin, H. (1931) Die Florideenordung Rhodymeniales. Lunds Universitets Arsskrift, N. F. Avdeling 2, 27 11: 1–48.

Kylin, H. (1956) Die Gattungen der Rhodophyceen. C.W.K. Gleerups Förlag, Lund, Sweden, 673 pp.

Lanfear, R., Calcott, B., Ho, S.Y.W. & Guindon, S. (2012) Partition Finder: combined selection of partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology and Evolution 29: 1695–1701.

Lawson, G.W. & John, D.M. (1987) The marine algae and coastal environment of tropical West Africa (second edition). Nova Hedwigia Beiheft 93: vi–415.

Le Gall, L. & Saunders, G.W. (2007) A nuclear phylogeny of the Florideophyceae (Rhodophyta) inferred from combined EF2, small subunit and large subunit ribosomal DNA: Establishing the new red algal subclass Corallinophycidae. Molecular Phylogenetics and Evolution 43: 1118–1130.

Le Gall, L., Dalen, J.L. & Saunders, G.W. (2008) Phylogenetic analyses of the red algal order Rhodymeniales support recognition of the Hymenocladiaceae fam. nov., Fryeellaceae fam. nov., and Neogastroclonium gen. nov. Journal of Phycology 44: 1556–1571.

Lin, S.-M, Fredericq, S. & Hommersand, M.H. (2001) Systematics of the Delesseriaceae (Ceramiales, Rhodophyta) based on LSU rDNA and rbcL sequences, including the Phycodryoideae, subfam. nov. Journal of Phycology 37: 881–99.

Lin, S.M., Fredericq, S. & Hommersand, M. H. (2012) Developmental, phylogenetic and systematic studies of Apoglossum and Paraglossum, Apoglosseae, trib. nov. (Delesseriaceae, Rhodophyta). European Journal of Phycology 47: 366–383.

Littler, D.S. & Littler, M.M. (2000) Caribbean Reef Plants. OffShore Graphics, Inc., Washington, D. C., 542 pp.

Lozada-Troche, C.A. & Ballantine, D.L. (2010) Observations on morphology and taxonomic affiliation of Coelothrix irregularis (Harv.) Børgesen (Rhodymeniales, Rhodophyta). Caribbean Journal of Science 46: 71–82.

Lozada-Troche, C., Ballantine, D.L. & Ruiz H. (2010) Cresia opalescens gen. et sp. nov. (Rhodymeniaceae, Rhodophyta) from Puerto Rico, Caribbean Sea. Cryptogamie Algologie 31: 293–303.

Norris, J.N. & Ballantine, D.L. (1995) Two new species of the red alga Chrysymenia J. Agardh (Rhodymeniales, Rhodymeniaceae) from the tropical western Atlantic. Proceedings of the Biological Society of Washington 108: 153–165.

Norris, R.E. (1989) Natalian Botryocladia (Rhodymeniales, Rhodophyceae) including description of a new long axis-forming species. Botanica Marina 32: 131–148.

Okamura, K. (1902) Nippon Sorui-meii [Book listing Japanese Algae]. Keigyosha, Tokyo, pp. i–vi, 1–276.

Okamura, K. (1936) Nippon Kaiso Shi [Descriptions of Japanese algae]. Uchida Rokakuho Publishing House, Tokyo, 9 + 6 + 964 + 11 pp.

Price, J.H., John, D.M. & Lawson, G.W. (1986) Seaweeds of the western coast of tropical Africa and adjacent islands: a critical assessment. IV. Rhodophyta (Florideae). 1. Genera A-F. Bulletin of the British Museum (Natural History) Botany 15: 1–122.

Rambaut, A. & Drummond, A. (2014) Tracer v1.6. Available from: (accessed 1 October 2017)

Rezak, R., Bright, T.J. & McGrail, D.W. (1985) Reefs and banks of the northwestern Gulf of Mexico: their geological, biological, and physical dynamics. Wiley, 259 pp.

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

Saunders, G.W. & Kraft, G.T. (1994) Small-subunit rRNA gene sequences from representatives of selected families of the Gigartinales and Rhodymeniales (Rhodophyta). 1.Evidence for the Plocamiales ord. nov. Canadian Journal of Botany 72: 1250–1263.

Saunders, G.W.I., Strachan, M. & Kraft, G.T. (1999) The families of the order Rhodymeniales (Rhodophyta): a molecular-systematic investigation with a description of Faucheaceae fam. nov. Phycologia 38: 23–40.

Schmidt, W.E., Gurgel, C.F.D. & Fredericq, S. (2016) Taxonomic transfer of the red algal genus Gloiosaccion to Chrysymenia (Rhodymeniaceae, Rhodymeniales), including the description of a new species, Chrysymenia pseudoventricosa, for the Gulf of Mexico. Phytotaxa 243: 54–70.

Schneider, C.W. & Lane, C.E. (2000) A new species of Botryocladia (Rhodymeniales, Rhodophya) from the Galápagos Islands. Cryptogamie Algologie 21: 167–175.

Schneider, C.W. & Lane, C.E. (2008) Notes on the marine algae of the Bermudas. 9. The genus Botryocladia (Rhodophyta, Rhodymeniaceae), including B. bermudana, B. exquisita and B. flookii spp. nov. Phycologia 47: 614–629.

Schneider, C.W. & Searles, R.B. (1991) Seaweeds of the Southeastern United States: Cape Hatteras to Cape Canaveral. Duke University Press, Durham & London, 553 pp.

Schnetter, R. (1978) Botryocladia monoica (Rhodymeniales, Rhodophyceae), a new species from the Caribbean coast of Colombia. Phycologia 17: 13–15.

Silva, P.C., Basson, P.W. & Moe, R.L. (1996) Catalogue of the benthic marine algae of the Indian Ocean. University of California Publications in Botany 79: 1–1259.

Silvestro, D. & Michalak, I. (2012) raxmlGUI: a graphical front-end for RAxML, Organisms Diversity & Evolution 12: 335–337.

Swofford, D.L. (2002) PAUP*. Phylogenetic analysis using parsimony (* and other methods). Version 4.1. Sinauer Associates. Sunderland, MA, USA.

Stamatakis, A. (2006) RAxML-VI-HPC: Maximum Likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics 22: 2688–2690.

Tamura, K., Stecher, G., Peterson, D., Filipski, A. & Kumar, S. (2013) MEGA6: Molecular Evolutionary Genetics Analysis version 6.0. Molecular Biology and Evolution 30: 2725–2729.

Taylor, W.R. (1960) Marine algae of the eastern tropical and subtropical coasts of the Americas. University of Michigan Press, Ann Arbor, 870 pp.

Wilkes, R.J., McIvor, L. & Guiry, M.D. (2006) Vegetative morphology and rbcL phylogeny of some members of the genera Botryocladia and Irvinea (Rhodymeniaceae, Rhodophyta). Phycologia 45: 481–94.

Wynne, M.J. (2005) Chrysymenia tigillis sp. nov. (Rhodymeniales, Rhodophyta) from the Sultanate of Oman, with a census of currently recognized species in the genus Chrysymenia. Phycological Research 53: 215–223.

Wynne, M.J. (2011) A checklist of benthic marine algae of the tropical and subtropical western Atlantic: third revision. Nova Hedwigia Beiheft 129: 1–166.

Yamada, Y. (1932) Notes on some Japanese algae, III. Journal of the Faculty of Science, Hokkaido Imperial University 1: 109–123, pls XXI–XXV.


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